What is that annoying buzzing sound?

After a period of dormancy on the blog, I finally have a little bit of time to write a few more posts. This is one of the last of the Churchill posts, and hopefully soon I'll be able to write about some new topics.

No recount of fieldwork in Churchill, Manitoba can be complete without some horror story of the bugs. While I was lucky during the 2009 season with abnormally cool temperatures, no such luck was had during the 2010 season. The mosquito density was just unreal. No amount of bug spray can protect you... I've been told that insect repellent does make a dent in the mosquitoes trying to bite you... instead of 5,000 mosquitoes trying to bite you, bug spray makes it so that a mere 3,000 are attacking you. In the end, the people who work in Churchill just don't find poisoning themselves with bug spray worth it. On the bright side, there are no diseases that mosquitoes carry in the far north!

During field work, we would have to be completely covered from head to toe. This involved wearing head nets, long sleeves, and, most importantly, gloves. Thin gloves were essential, especially when we had to take field notes. In addition, when dealing with blackflies, it was extremely important to tuck pants into socks, gloves into sleeves, and shirts into pants, since blackflies crawl into tight spaces and will find their way into the tiniest hole to bite you. When all covered and prepared, I usually avoided mosquito bites in the field completely. I usually ended up getting more bites in the Study Centre or in the car.

Now, words can only do so much to describe the bug density. For the rest, I will let my pictures show the horror.

A large flock of mosquitoes.

Gloves are important.

Mosquito carnage. In the top image, I got 72 in one swat of my hand. I can't remember how many we got in our field notebook, but it was well over 100.

Yours truly, doing veg surveys, with a small cloud of mosquitoes around my head.

A truly scary cloud of mosquitoes. Click on the image to zoom in to see all the mosquitoes that are a bit farther away.

The next few photos are from Jay McGowan, who I spent 2009 field season with. His pictures truly capture the insect abundance.

Mosquitoes like dark colors. Zoom in on this picture to see how many are really there. It's unreal. (photo by Jay McGowan)

I think this may be one of the signs of the Apocalypse... "and the Lord sent forth a cloud, and the sky was darkened with mosquito" (photo by Jay McGowan)

The following video is from the 2010 field season on one of the more buggy nights. Listen with sound...

Bird species discovery odds and ends

Here are a couple more neat graphs about the rate of discovery of new bird species that I couldn't fit well into my previous two posts about the subject (10,000 birds? Part 1 and Part 2).

In Part 1 I briefly introduced the species accumulation curve as a means of comparison of rates of discovery between different groups. Here, I wanted to parse apart any broad-scale patterns in the bird data. I started by splitting the birds into their two broadest groupings - passerines (5927 species) and non-passerines (4100 species) - and then I split passerines again into two more broad classes - oscines (4668 species) and suboscines (1259 species). These new species accumulation curves are plotted against the overall bird curve (in black):

(click to enlarge)

The non-passerine curve falls generally to the left of the passerines and birds overall, indicating that non-passerines were generally described earlier than passerines and fewer non-passerines have remained undescribed until recently. The passerine, oscine, and suboscine curves are generally similar in slope and position, although suboscines lag in the last 100 years or so. The suboscine curve is also the only one with a noticeable uptick - a bunch of new species described - in the last few decades.

Those who know me when I get into projects like this know I like to take things to ridiculous excess. In this case, I decided to investigate family-level patterns of description. To get fairly smooth accumulation curves requires a fairly large number of taxa, so I picked the bird families recognized by Birdlife that contain more than 150 species. These twenty families are:

Family name	Species
Tyrannidae	414
Psittacidae	374
Trochilidae	337
Timaliidae	326
Columbidae	318
Emberizidae	313
Sylviidae	293
Muscicapidae	286
Thraupidae	256
Furnariidae	241
Accipitridae	238
Picidae	218
Thamnophilidae	218
Strigidae	186
Phasianidae	181
Meliphagidae	177
Fringillidae	177
Turdidae	173
Anatidae	164
Rallidae	156

And their species accumulation curves, plotted against the all-bird curve in black:

(click to enlarge)

I don't expect anyone to actually tease apart that rainbow coalition of squiggles in detail, especially because they form such a tight column of similarly-sloped curves. That in itself is a pretty neat result - the family-level differences aren't as noticeable as I thought they would be, and teasing them apart in detail is fairly pointless. There are, however, two curves that stand apart from the rest.

On the far left in pale blue is Anatidae. 50% of the currently known ducks were described by 1800, decades before any other family, and 90% were described by 1870. Ducks are big, obvious birds with common interaction with humans (in the form of hunting) and often bold, distinct male plumages in each species. It is thus not surprising that ducks were described comparatively early in history and very few new species have been described in the last 100 years.

In contrast, on the right side of the curves, the pale reddish curve for Strigidae falls below the others in the last 100 years or so. The slope is pretty similar to the other curves for this time period, so I don't think the overall rate of discovery in owls is much lower. Instead, the curve is displaced lower by the flush of new species described in the last few decades - remember in my last post that owls had the highest number of new species described of any family since 1942.

That's all I have for now - have fun staring at those colorful squiggles.

10000 birds? Part 2

In Part 1, I looked at the history of bird species discovery, and found that new bird species have been described at a fairly steady rate of an average 4.9 species per year since the 1940s. Here, I'll review what species have been discovered since the 1940s, where they are found, and why it took so long to find them.

From 1942 until 2008, 330 species encompassing 76 families and 213 genera were described:

(note: I accidently included 1 2009 datapoint, but 2009 is incomplete and more than 1 were described that year)

Among the top families, five (Tyrannidae - tyrant flycatchers, Thamnophilidae -antbirds, Furnariidae - ovenbirds, Rhinocryptidae - tapaculos, Formicariidae - anthrushs) represent the diverse suboscine lineages of South America. Two other top families (Sylviidae - old world warbles and Timaliidae - babblers) are similarly diverse lineages from the old world (I don't think Birdlife carves up these families into smaller groups as other checklists do in the last few years). My gut impression before seeing this data was that these two broad groups of birds would be the leaders for multiple reasons - they compose many of the species I could remember seeing described in the last few years, they contain many cryptic groups and are receiving active taxonomic treatment with new input from DNA and song, and they are found in the tropical areas of the world still receiving new exploration and documentation of fauna. However, the winner in the family category by a large margin was a surprise for me: Strigidae (owls). It makes sense that cryptic nocturnal birds would contain a lot of previously undocumented diversity, but I was surprised at the scale. The top genera track the top families as expected (Glaucidium - Pygmy-Owls, Otus - Scops-Owls, Scytalopus - Tapaculos, Grallaria - Antpittas, Phylloscartes - Tyrannulets, etc).

There are a myriad of historical and unquantifiable reasons that bird species persist until the present day undiscovered or unrecognized by science. However, I made several simple, straightforward predictions about recently discovered species that can be quantified and tested:

1) New species are likely to be found in underexplored tropical countries
2) New species are likely to be single-country endemics
3) New species are likely to have small ranges
4) New species are likely to have small populations and thus more likely to be endangered

Using Birdlife International's online database, I was able to extract relevant bits of information regarding these new species - where they are found, whether they are country endemics, their estimated range size, and their Birdlife conservation status. I tested my predictions by comparing the species described from 1942-2008 with random samples of species described earlier in history: 100 species described by Linnaeus in 1758 and 200 described in the 1840's-1860's.

1) New species are likely to be found in underexplored tropical countries

Here is a summary of where the new species (1942-2008) are found. The first column is overall country listings, which sums to well over 330 because of species ranging across many countries. The second column of results is for single-country endemic species.

As expected, the South American and Southeast Asian tropics top both lists, with Peru and Brazil leading both categories by a good margin.

2) New species are likely to be single-country endemics

I didn't tally the country-by-country data for my 1758 and 1800's sample groups, because so many of them span a huge range of countries. This difference is apparent when you compare the proportion of single-country endemics for each category:

6% of species described in 1758 are found in a single country
24% of species described in the 1840s-1860s are found in a single country
72% of species describe 1942-2008 are found within a single country

This confirms that recently described species are more likely to be restricted to within a single country, and are thus easier to overlook, than those species described earlier in history.

3) New species are likely to have small ranges

I binned the Birdlife range size data by order of magnitude for each of the three time periods and calculated the proportion of the sample in each:

(click to enlarge)

The species described by Linnaeus in 1758 (blue bars) are overwhelmingly species with huge ranges in the millions of square kilometers. Species described in the mid-1800's still have ranges mostly in the hundreds of thousands to millions of square kilometers, but there is a wider distribution of species among the size classes.

For new species described 1942-2008, the distribution among size classes is much wider than in the other time periods. Less than 4% of these species have ranges in the millions of square kilometers - these include species like Cryptic Forest Falcon (Micrastur mintoni) and Amazonian Pygmy-Owl (Glaucidium hardyi) which have large Amazonian ranges. The largest proportion of new species have ranges sizes that are fairly small - 1000s to tens of thousands of square kilometers.

The most stunning category is for range sizes of less than 100 square kilometers - an area only 5 times bigger than my hometown of North Tonawanda, NY. 12.5% of new species (32 total) fall into this category, with some absurdly small known ranges. Four species have a known range of less than ten square kilometers - Munchique Wood-Wren (Henicorhina negreti, 8 sq. km), Poo-uli (Melamprosops phaeosoma, 3 sq. km), Calayan Rail (Gallirallus calayanensis, 2 sq. km), and Bugun Liocichla (Liocichla bugunorum, 2 sq. km). Words fail me in trying to describe how mind-boggling, fascinating yet even heart-breaking it is to see species with ranges like this. The only species with ranges this small in the other categories are a fruit-dove and a honeyeater described in the mid 1800s from tiny South Pacific islands with range sizes around 25-50 sq. km.

4) New species are likely to have small populations and thus are more likely to be endangered

I wasn't able to easily pull data on population sizes for a large range of species from the Birdlife database, so instead I used a proxy - Birdlife conservation status. I calculated the proportion of species from each time period that fall into the six Birdlife categories:

DD = Data Deficient
LC = Least Concern
NT = Near Threatened
VU = Vulnerable
EN = Endangered
CR = Critically Endangered

(click to enlarge)

Just as the 1758 and mid-1800s samples were overwhelming skewed towards species with large range sizes, these categories are both over 80% composed of species ranked Least Concern. Only one of the 1758 species (Bald Ibis, Geronticus eremita) and one of the 1800s species (Cuban Kite, Chondrohierax wilsonii) are recognized as Critically Endangered. A plurality of the new species are Least Concern, but there is a very wide distribution of rankings, with 8.5% (28 species) being Critically Endangered.




Well, that about covers it. If you weren't keeping a running tally, all of my predictions were validated by the data! While I love playing with this data and learning more about the patterns of discovery of new species, I realized in the end that these findings are rather disturbing. A depressingly large proportion of bird species that have remained unknown to science until recently have very tiny ranges and are endangered. In this era of exceptional environmental destruction, how many more species like these have slipped away into extinction unnoticed? That is a number we can never know.

10,000 Birds?

"You think you know every bird in the world, and then someone shows you a Curl-crested Aracari"

David Quammen, The Song of the Dodo

An oft-cited figure for the diversity of birds in the world is around 10,000 living species, a reasonable rule of thumb although estimates vary - Birdlife International currently recognizes 10,027 while the IOC list is currently at 10,396. Duncan Wright says the number should be closer to 12,000 if recently extinct birds are included (and he also coincidentally begins with a David Quammen quote). Pinning down an exact number is an impossibility because differing opinions on species limits in species and subspecies complexes create a near-infinite number of list permutations. Pinning down a number is also impossible because the bar keeps getting raised by the new bird species discovered every year.

New species of bird often receive a small splash of press when discovered or described, because birds are regarded as an exceptionally well described group of vertebrates and we've long reached the point where new birds are few in number. Additionally, the hunt for new species has been romanticized in such good reads as A Parrot Without A Name or this more recent Birding article (pdf) by Joseph Tobias. Tobias quotes Mayr (1946) as proclaiming the general end of new bird discovery - “I doubt that in the entire world even as many as 100 new species remain to be discovered” - but states that a yearly trickle of new species have been described since then. Just how many new birds are discovered every year? How much has the trickle of new birds declined? How many new birds are left to discover?

I can't answer the last question, but I can find data for the first two. It is a logistical nightmare to track down this data by finding every new species description in the primary literature. Instead, I picked a world checklist to find what I needed. The formal way to list species in literature includes a citation for the description of the species, such as Puffinus griseus (Gmelin 1789). By using these dates, you can quickly extract the year by year data for species descriptions. This method also only counts newly described species, not splits through the elevation of previously recognized subspecies.

The only quickly downloadable checklist I found with data like this is Birdlife International's checklist, so I used their most recent version (v3, June 2010) and plotted out the data. Here is the number of new bird species described each year, beginning in 1758 with Linnaeus and ending in 2008 (the Birdlife checklist is incomplete for new species in 2009 and 2010):

(click to enlarge)

The early decades show relative inactivity in bird taxonomy punctuated by massive monographs (like the 400+ that Linnaeus described in one work). The golden age of bird taxonomy peaks in the mid-1800's, with several decades of 50+ species described every year. The number of new birds described then generally declines to the present day.

Another way to present this data - the most useful for comparison - is to create a species accumulation curve. This represents, for each year, what percentage of the total number of known species was described at that point. This allows you to examine what those small numbers of new species each year mean in the context of the total number of known bird species. The slope of the curve is a proxy for the rate of discovery of new species, and you can easily eyeball the differences in slope representing different rates of species description over time.

Here is the species accumulation curve for birds:

(click to enlarge)

This represents a typical curve for a well-described fauna - the middle range is steepest, and the curve plateaus as new species become increasing hard to find. Compare the curve for birds with that of a very different fauna, another favorite of mine, geckos:

(click to enlarge)

In stark contrast to birds, the rate of discovery of geckos has only accelerated through time, and the last few years have seen the highest yearly rates of gecko discovery ever - but that's another post. The point is that geckos are a representative of an under-described group with lots of new species to be found and taxonomic description left to be done. Birds are comparatively well-described and have relatively little left to be found.

We can estimate how little is left to be found by studying the decline in new species found. So how about that decline? Zooming in to the last 120 years, we get this:

(click to enlarge)

At about the time Mayr proclaimed the end of bird discovery in the 1940s, the overall rate of description of new birds had been trending steeply downward for decades. After the rate reached rock bottom - 1941 is the last year with more than ten bird species described - there is indeed a steady trickle of new species every year, averaging 4.9 species per year and varying from zero (in 1954 and 1978) to ten (1960, 1974, 1997) new birds yearly.

However, this trickle does not seem to show any sign of stopping, even decades after Mayr's prediction. In fact, if you take just the data from 1942-2008, look at the general trend line:

(click to enlarge)

That's right, that is a (very weakly supported) POSITIVE trend line for the discovery of new species in the last seven decades. Given the overall wide variance in discovery from year to year in these decades, I double-checked this trend my making a moving ten-year average of new birds per year:

(click to enlarge)

This is a really fascinating result! Despite the fact that the overall description rate of birds has plummeted and won't ever get back up to the rate it was 100 years ago, discovery of new birds has stabilized and shows no signs of slowing since the 1940's. That means that, while the species accumulation curve has plateaued, it is not exactly drifting towards zero and the total number of remaining bird species can't be extrapolated. Who knows what is left to find?

In part 2 of this post, I'll outline what new species have been discovered in the last few decades, where they are being found, and some reasons for their late discovery. Stay tuned.