We have gained many ecological and evolutionary insights from studying variation in DNA markers, from resolving the very base of the tree of life (genealogical affinities of bacteria, archaebacteria/extremophiles, eukaryotes), through overturning received truths about mating systems of birds (most are not in fact genetically monogamous), to quantifying impacts of human activities on connectivity of populations of species of conservation concern. Among the many revelations that come from such DNA studies are those from phenotypically cryptic taxa whose appearances often mask deep phyletic diversity. Indeed an increasing number of studies shows that myriad, traditionally-regarded ‘species’ are in fact complexes of separate, reproductively-isolated species. DNA studies have revealed such cryptic species in many groups, including mammals (Ceballosa & Ehrlich. 2009), birds (e.g. Lohman et al. 2010), amphibians (e.g. Elmer et al. 2007), and insects (e.g. Hebert et al. 2004). Many examples of cryptic diversity come from the tropics, but here are also some intriguing examples from higher latitudes like ours, with implications not only for understanding of evolutionary affinities of taxa in question, but also for their geographical distributions and the forces that have shaped them.
The trilling chorus frogs (a distinct lineage or clade within the treefrog genus Pseudacris) comprise one such group. This clade, distributed broadly across eastern North America, includes at least nine species: the mountain chorus frog (P. brachyphona), Brimley’s chorus frog (P. brimleyi), spotted chorus frog (P. clarkia), Cajun chorus frog (P. fouquettei), New Jersey chorus frog (P. kalmi), upland chorus frog (P. feriarum), southern chorus frog (P. nigrita), boreal chorus frog (P. maculata) and western chorus frog (P. triserieta) (Moriarty & Cannatella 2004).
Two of these species occur in Ontario, P. maculata and P. triserieta. The two are very similar in appearance – both are small (generally < 3.5 cms in snout-vent length), with smooth skin, and a dorsum varying in colour from brown to greenish-gray, and a dark stripe through the eye and longitudinal markings on the dorsum. The calls too are very similar being comprised of a trill that is often likened to running one’s fingers along a plastic comb. The boreal chorus frog was until recently considered to be distributed from northwestern Ontario to Alberta and north to the NWT, also being found in the USA in the Midwest south to Arizona and New Mexico. The western chorus frog was thought to range from southern Quebec and Ontario/northern New York state west to South Dakota, and south to the states of Kansas and Oklahoma (Harding 1997). In southern Ontario until recently there were considered to be two regional populations of P. triserieta: a “Carolinian population” found south and west of Toronto, and a Great Lakes–St. Lawrence population found east and north of Toronto, with the latter considered as ‘Threatened’ under the Canadian Species at Risk Act.
That’s the old view. Mitochondrial DNA evidence suggests that the Great Lakes–St. Lawrence population which was classified as P. triserieta is not in fact western chorus frog at all, but rather is a disjunct population of boreal chorus frog (Lemmon et al. 2007a,b, Rogic et al. 2015). Playbacks by Rogic et al. (2015) seem to affirm this, with eastern Ontario and western Quebec chorus frogs responding to previously recorded calls of P. maculata and not P. triserieta.
All of this has interesting implications for 1. Conservation (Is this western boreal chorus frog population genetically distinct and thus does it merit conservation priority?), 2. Biogeography (How did the species become disjunct and what paths of re-colonization did these distinct populations use?), and 3. Understanding the nature of species (these trilling chorus frogs are cryptic to us, but clearly they can tell each other apart – it is in the domain of mate recognition system and acoustics that the species differences are clear). As always there’s lots more work that can be done, not least of which is more finely mapping genetic diversity across the entire boreal chorus frog distribution.
Ceballosa, G. & P.R. Ehrlich. 2009. Discoveries of new mammal species and their implications for conservation and ecosystem services. Proc. Natl. Acad. Sci. USA 106: 3841–3846.
Elmer, K.R., J.A. Davila & S.C. Lougheed. 2007. Cryptic diversity, deep divergence, and Pleistocene expansion in an upper Amazonian frog, Eleutherodactylus ockendeni. BMC Evol. Biol. 2007, 7:247.
Harding, J. 1997. Amphibians and Reptiles of the Great Lakes Region. Univ. Michigan Press. Ann Arbor, MI.
Hebert, P.D.N., E.H. Penton, J.M. Burns, D.H. Janzen & W. Hallwachs. 2004. Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator. Proc. Natl. Acad. Sci. USA 101: 14812–14817.
Lemmon, E.M., A.R. Lemmon, J.T. Collins, J.A. Lee-Yaw & D.C. Cannatella. 2007a. Phylogeny-based delimitation of species boundaries and contact zones in the trilling chorus frogs (Pseudacris). Mol. Phylogenet. Evol. 44:1068–1082.
Lemmon, E.M., A.R. Lemmon & D. C. Cannatella. 2007b. Geological and climatic forces driving speciation in the continental distributed trilling chorus frogs (Pseudacris). Evolution 61: 2086–2103.
Lohman, D.J., K.K. Ingram, D.M. Prawiradilaga, K.Winker, F.H. Sheldon, R.G. Moyle, P.K.L. Ng, P.S. Ong, L.K. Wang, T.M. Braile, D. Astuti & R. Meier. 2010. Cryptic genetic diversity in “widespread” Southeast Asian bird species suggests that Philippine avian endemism is gravely underestimated. Biol. Conserv. 143: 1885-1890.
Moriarty, E.C. and D.C. Cannatella. 2004. Phylogenetic relationships of the North American chorus frogs (Pseudacris: Hylidae). Mol. Phylogenet. Evol. 30: 409–420
Rogic, A., N. Tessier, S. Noël, A. Gendron, A. Branchaud & F0J. Lapointe. 2015. A “trilling” case of mistaken identity: Call playbacks and mitochondrial DNA identify chorus frogs in Southern Québec (Canada) as Pseudacris maculata and not P. triseriata. Herp. Rev. 46: 1-7.
(unless otherwise credited, photos by Art Goldsmith)
Field Course 2015: Effects of Human Development on Aquatic Environments and Biodiversity in Canada and China
World-Wide Fish Physiology and Ecology with Professor Wang AND Views from QUBS
Professor Yuxiang Wang and Fish Facts
On July 30, 2015, Professor Yuxiang Wang presented his lecture on Fish Biodiversity/Physiology and Conservation. The slide presentation roamed the globe through Yuxiang’s eyes, to many of the most diverse and least diverse ecosystems. His breadth and depth of observation, analysis and research kept the students and me in maximum learning mode.
The assembled class put down their cell phones and looked their very best for this photo, as they awaited Professor Wang.
The basics and fish facts are required to understand the more detailed knowledge imparted on us later in the session.
Any excellent introductory course will inform participants about the big picture. The slides above and below answer the questions: How many kinds of fish are there and where did they come from? Fish are indeed vertebrates, animals with backbones, just like us and the other mammals, birds, reptiles and amphibians. And there are lots and lots of different kinds of fish.
Canada’s record on marine and freshwater conservation isn’t the brightest, with less than 1% of our oceans under any kind of protection, and that protection is very minimal, as resource industries still take priority even in our marine parks. Australia, for contrast, has managed to protect 38% of its surrounding marine space. When I saw Professor Wang’s slide above, it reminded me of the quest to save diversity among species, since diversity is the underpinning of a healthy biosphere.
I recall visiting Biscayne National Park in Florida, USA, where one of the pamphlets proclaimed that there were more fish species in this national marine park than all the vertebrates in Colorado put together. These fish species require a healthy shoreline wetland environment and unpolluted water to persist. Biscayne National Park also has a large coral reef, another important and stressed element for marine ecology globally.
Evolutionary biology and the origins of fish are subjects that require large volumes to achieve basic understanding. Yiuxang’s slide is a great summary. Note the exclusions from generalities about fish (lamprey and hagfish). The lamprey is of special interest for me, as it was one of the first alien species to reach our Great Lakes and St. Lawrence River, threatening an already stressed fishery. Having not focused on this species in decades, Yiuxiang provided a good summary of recent developments regarding attempts to control the Sea Lamprey, (Petromyzon marinus). Of special note, the lamprey is native to the North Atlantic. It is anadromous (more later) and could reach the Gulf of St. Lawrence. The St. Lawrence Seaway project of the1950s provided convenient routes around rapids and falls, yielding a fresh supply of fish to larval lamprey throughout the St. Lawrence and Great Lakes.
Hagfish and Lampreys
The Jawless fish kicked (hard to do without feet!) things off, specifically, Hagfish and Lampreys.
The 60 species of Hagfish with rudimentary eyes live in hypoxic habitats (low oxygen) with little light (i.e., they may live in ocean depths). Here are a few more facts about Hagfish from U.C. Berkeley:
“The adjective which best describes the Myxini is “Lovecraftian”. Hagfish are long, slender and pinkish, and are best known for the large quantities of sticky slime which they produce. Hagfish have three accessory hearts, no cerebrum or cerebellum, no jaws or stomach, and will “sneeze” when their nostrils clog with their own slime. They are found in cold ocean waters of both hemispheres, scavenging dead and dying fish but also preying on small invertebrates.
Hagfish are almost blind, but have well developed senses of touch and smell. They have four pairs of sensing tentacles arranged around their mouth. The mouth lacks jaws, but a Hagfish is equipped with two pairs of tooth-like rasps on the top of a tongue-like projection. As this tongue is pulled back into the Hagfish’s mouth, the pairs of rasps pinch together. This bite is used to tear into the flesh of dead and dying fish which have sunk to the muddy ocean bottom, or in catching and eating marine invertebrates. By far, the largest part of their diet is polychaete worms. Due to their slow metabolism, Hagfish may go for up to seven months without eating any food.”
I bet the creature in the “Alien” movies was based on the hagfish and lamprey.
Their “vertebrate” status has also become compromised through adaptive evolution, as their spinal column is vestigial (no vertebrae) with a partial cranium. There are hagfish in the Gulf Islands off Vancouver.
Instead of tooth-like Hagfish rasps, the lamprey’s mouth evolved suctorial disc teeth. The front end of a lamprey is a sight, looking more like a sanding disk with its circular array of inward pointing teeth. They do not have the paired fins of most other fish. Indeed, like the hagfish, lampreys attach themselves to a fish, rasping away flesh using the “buzzsaw” mouth. Ouch! They are also anadromous. Like Atlantic Salmon, the adults spawn and develop in freshwater and then return to saltwater to live. Larvae live buried in freshwater systems. Biologists applied their knowledge of the lamprey to control it. Lamprey use chemo-sensing, rather than sight, to manoeuvre through their habitat. Like many insects, they use pheromones to find each other for breeding. Dr. Lee, at Michigan State University, used this knowledge to attract males, which were then sterilized. Returning to the environment to mate, these males, of course, produce no offspring, reducing (not eliminating) populations. This application of biology is much more refined, effective, economical and environmentally benign than the first approach to control: chemicals, which kill much more than the target species. The photo above, from Yuxiang’s presentation shows a lamprey attached to and feeding on a Lake Trout. The lowest photo in the slide shows the mouth of the lamprey.
Now that we have dealt with the exceptions, let’s survey the rule—the bony fish.
Cartilaginous and Bony Fish
The sharks and rays are classified in a separate group (Class Chondrichhyes), which are considered to be older than other jawed fish on the evolutionary scale. The Holocephali (Ratfish and cousins) are less well-known deep-water living marine fish.
The bony fishes (Class Osteichthyes), are the rest of the fish species and the most familiar to us. Avid fishers know well the species that respond to our lures and bait. There are so many more. One of the more illustrious species, the Coelacanth, a living fossil (It was thought to have become extinct 80 million years ago. Oh well, science does learn from mistakes!) was featured prominently in my McGill University education in the late 1960s.
Read this fascinating story from the Washington Post, and this other article from the Australia Museum:
The discovery of the Coelacanth has filled in answers to evolutionary questions about the evolution of bony fish and the tetrapods (those of us with legs).
This brief background led Yuxiang into the meat and potatoes (or fish and chips?) portion of his presentation: adaptive and ecological physiology of fish.
Teleosts are any member of a large and extremely diverse group of ray-finned fish. Along with the chondrosteans and the holosteans, they are one of the three major subdivisions of the class Actinopterygii, the most advanced of the bony fishes. The teleosts include virtually all the world’s important sport and commercial fishes, as well as a much larger number of lesser-known species. When the average person thinks about fish, he/she usually has a teleost in mind.
The slide above describes how fish have adapted structurally and functionally to their environment. The following terms are important to understanding fish physiology in some of the more extreme habitats studied by Professor Wang. The slide below maps out typical teleost body parts.
Ammonotelic refers to a fish that excrete nitrogenous waste derived from amino-acid catabolism in the form of ammonia.
Poikilotherm refers to an organism that cannot regulate its body temperature except by behavioural means, such as basking or burrowing.
We hear daily about increasing pressure on our marine and freshwater species. Fish are staples of the human diet. Therefore, conservation would logically move to the top of our global “to do” list. That isn’t the case, and research like that of Professor Wang’s aids those of us with the motivation to act and provides us with the evidence needed for change. Of course, the Queen’s University Biological Station (QUBS) is a key resource for freshwater fisheries research. Researchers come from all over North America to study the fish in Lake Opinicon and other surrounding lakes. There are also lakes wholly contained within the QUBS properties, making them ideal for whole lake research. Lake Opinicon’s shore houses the QUBS facilities. It is quite a diverse lake. In fact it is a bit more diverse than the list given in the slide below. The two fish pictured in the slide are two other very common Centrarchidae, the Blue Gill (Lepomis macrochirus), and Pumpkinseed (Lepomis gibbosus) Sunfish. These two fish are often the first fish a child fisher may catch in our area. There is a Centrarchidae missing: the Black Crappie, (Pomoxis nigromaculatus) seen in the photo beside “Fish in Lake Opinicon”. Those are Gregory Bulte’s hands briefly showing the fish to his class this summer at QUBS. Look for another edition of this blog featuring Grégory Bulté’s Ecology Field Course.
Onward to Professor Wang’s research, which focuses on fish physiology and ecology. Back in my student days, the term physiology caused me to shudder and run. If only Yuxiang Wang’s approach had been available back then. He made a clear and easy-to-understand connection between physiology and ecology. Also, he makes a good case for this kind of study to better understand fish which are important to the health and well being of local people. I especially appreciate the non-laboratory approaches that Wang takes. He gets to know and to study his fish on site in some of the more interesting places on Earth.
Who wouldn’t call Amazon waters “amazing”? Some of Professor Wang’s work is done at the confluence of two main Amazon tributaries— the Rio Solimoes and the Rio Negro— at Manaus, Brazil. Professor Wang is telling us, by pointing at the water chemistry slide, to look at the amazing water chemistry differences between the two.
The Rio Negro’s black waters are a result of their origin in the mineral poor tropical rainforests of northwestern Brazil. Alternatively, the Rio Solimoes originates in the Andes and covers a great deal more distance than the Rio Negro. Therefore, its waters are mineral rich. The Rio Negro is acidic, much like our boreal rivers that carry heavy loads of carbonic acid, while Rio Solimoes water is almost neutral. As its name implies, the waters of the Rio Negro are very black when seen from aerial or satellite views, whereas those of the Rio Solimoes are muddy and creamy looking from the same views. Even after the rivers come together, you can see these colour differences far downstream. Of course these differences mean that fish living in each river are adapting to these very different conditions. Therefore, this is an ideal spot for comparative fish physiology.
From the category “I did not KNOW that!” Yuxiang tells us that the intense flow of the Amazon River pushes its fresh water 300 kms out to sea in the South Atlantic. Ships crossing this band of fresh water would sink if they were too heavily laden, due to the drop in buoyancy crossing from salt to Amazon River water.
The Pirarucu (Arapaima gigas), shown above, grows to enormous sizes in the challenging Amazon waters. Like most fish, (tuna and billfish are exceptions), Pirarucu are poikilotherms, that is their body temperatures change with environmental temperature changes. Since water has a high heat capacity, much higher than air, poikilotherms have less of a challenge in water than terrestrial vertebrates. This gigantic Amazon fish adapts in many ways. In the very warm oxygen-poor waters of the Amazon, Pirarucu breathe air using an adapted swim bladder. Another extraordinary adaptation is the Pirarucu kidney. Most animals need a way to excrete nitrogen, a necessary by-product of body waste management. Birds, reptiles and insects excrete uric acid, and mammals excrete urea. These latter biochemicals avoid the toxicity of the simpler nitrogen compound, ammonia. Therefore, it is astonishing to learn that the Pirarucu is ammonotelic, excreting ammonia. The warm Amazon waters do lack oxygen, and daily oxygen and temperature fluctuations may also be great.
The Oscar (Astronotus oscellatus), pictured below, is highly adapted to avoiding hypercapnia, excessive carbon dioxide in the bloodstream, typically caused by inadequate respiration.
Just for fun: did you know that teleosts make up such a large part of the oceans’ biomass that their “poop” maintains the pH of the waters, regulating acid-base levels.
To China we go…. Professor Wang has conducted ongoing research at the remarkable Lake Qinghai, a very large lake on the cold, dry north central plain of China. Recent socioeconomic developments and internal migration have changed the local human population culture. The original inhabitants have great respect for the lake and its formally teeming fish species, the Naked Carp (Gymnocypris przewalski). The newly arrived inhabitants have decided to develop irrigation-based agriculture in the lands around the lake, which has resulted in a 10-12 cm drop per year in lake levels. The pH of the lake is 9.4 (very basic), and it has a high Magnesium and low Calcium concentration. When lake volume decreases, salinity increases, putting the animals adapted to the natural pH at risk. You may read about the lake’s story in the work cited in the slide below (Wood et al.) and by following up with Professor Wang’s research.
The next slide cites “Wood et al” and their work on how the carp have adapted to the rapid (only 50 years) decline in Lake Qinghai water volumes and increased salinity. Wang’s work has shown that much needs to be done to protect the lake and its biological community, as imbalances threaten its most vulnerable rare species.
The students also had many field opportunities during their two weeks in Canada. They toured the Thousand Islands and the Kingston sewage treatment plant. They went to Parliament Hill in Ottawa and then, very central to any Canadian biologist’s bucket list, visited the collections of the Canadian Museum of Nature in Aylmer, Quebec. Thanks to Team Round-headed Apple Borer and to Mark Szenteczki for the following photos:
Thanks to the students and Mark, and these two course creators,producers, directors and stars. We will see you in China for this course next year!
Tree Swallows research is the next topic, and look for stories and photos featuring Professor Raleigh Robertson, the long-time Director of QUBS, who started Tree Swallow research at Queen’s in the 1970s. Also, some photos of QUBS will be featured.
(unless otherwise credited, photos by Art Goldsmith)
Effects of Human Development on Aquatic Environments and Biodiversity in Canada and China Field Course 2015
Chapter 2 – Student Seminars
Invasive Species Seminar
Environmental and Ecological Impacts of Dams by Team Scrambled-Egg Slime Mold
Seeing this topic connects me to relevant strong and vivid personal memories. Since World War 2, so much of human enterprise has been about dams, principally those built to satisfy our gluttonous appetite for cheap energy. In the 1970s, I worked with Professor John Spence of McGill University, who took on the role as Science Advisor for the First Nations’ court battle against Hydro Quebec’s massive James Bay development (1972). Those dams are now built, and the environmental and social consequences well known.
Therefore, it was with more than a little interest that I entered into this discussion with these four students.
Canada has a large share of hydro development mega-projects, including James Bay (Quebec), Churchill Falls (Newfoundland and Labrador), St. Lawrence–Great Lakes (Ontario), Churchill–Nelson (Manitoba), plus so many more megawatts of capacity on most of our streams in the populated south of the country. When you add water supply, navigation and irrigation dams, finding natural rapids and falls in southern Canada is now a challenge! Some species, like the Rapids Clubtail dragonfly (Gomphus quadricolor), which is dependent on southern rapids for habitat, are now rare and endangered. Hydrological and stream hydraulics changes cause an array of undesirable ecological effects.
Although no southern projects have recently caused major social disruption in Canada, the same is not true for China. In Canada, only the Eisenhower Dam project, which created Lake St. Lawrence, comes to mind. In that 1950s case, three villages were flooded. Most of the homes and businesses were moved to a larger planned village nearby (Ingleside, Ontario) before flooding. Historical buildings wound up at a new interpretive historical village (Upper Canada Village, Morrisburg, Ontario).
In China, when we hear that 1.3 million people lost their homes to the Three Gorges Project, the scale boggles the mind. Already threatened ecosystems and their species are now contending with new development pressures. Just opened in 2003, this dam across the Yangtze River is associated with the largest capacity power station on our planet. The reservoir is a 1,000-square kilometre lake which reached its “final height” in 2010.
One can only imagine the ecological and socioeconomic consequences. But why do that, when, like our four team members, you may easily read the facts yourself.
Oil Spills by Team White-winged Scoter (Melanitta deglandi)
Team Members: Chang Leo, Mengxi Wu, Anyi Tang, Manreet Kaler
My time with this group was short. I talked with Chang before the other three team members arrived. He started by telling me about the different kinds of oil and oil products which are transported and spilled:
1. very light
Of the four types, very light is the most difficult environmentally due to its volatility and high toxicity. We all know well the results of offshore drilling infrastructure collapse (BP’s Gulf of Mexico drilling platform) and transportation accidents (pipeline breakages and shipping accidents, the Exxon Valdez in Alaska being one of the best known and most destructive).
The group also reported on one of the activities on the course blog. On August 3rd, the students participated in a fish seining learning exercise. Seining is one of the most straightforward ways to sample the population of fish along a shoreline. Partners spread out after extending a net to its full length and attaching it to the appropriate foot. The two people at each end then move out, so as to create a pocket with the net. They then envelop the fish by moving toward each other, closing off the pocket. Fish are transferred to buckets for identification, counting, and data collection (age, sex, condition, etc.). This team caught 158 fish, 7 species.
Aquaculture in the Modern Era by Team Black-shouldered Spinyleg (Dromogomphus spinosus)
Huck Nelson is from PEI and feels a vested interest, like so many young Maritime biologists, in the burgeoning aquaculture industry. His hopes and concerns were echoed by the Chinese members of this team.
Upon meeting, we immediately entered into an animated discussion about the origins of aquaculture. The team found reference to British Columbia First Nations’ clam farming as far back as 5,000 years. Evidence of fresh water fish farming in Egypt would be concurrent, and carp in china may reach back as far as 4,500 years. So there is a long-term human penchant for stewardship of aquatic animals in addition to terrestrial animals.
At Queen’s, the Bruce Tufts’ lab in Fish Physiology and Fisheries Biology is investigating the effects of different applications on aquaculture fish through their fish physiology expertise. For example, the team learned that a flow-through system is 200 times less efficient than a bio-filtered re-circulation system.
In terms of environmental impact, aquaculture is having a variety of undesirable outcomes; the most concerning being the weakening of wild populations, local pollution (flow-through) and the focusing on mono cultures.
Chinese aquaculture dwarfs our own, as 70% of global aquaculture production is Chinese. Their products do show up in our stores. Asian shrimp production (India, Vietnam, Malaysia, Thailand and China) has become big business, as most of the large volumes of shrimp now sold to North America come from that region.
I learned so much from this and subsequent discussions. For me, that is the big payback. Learning in such an enriched environment is a privilege. Thanks, Xuewei, for the Chinese lesson! “Yu” means fish; “tsing” is please; and Beijing has 24 million people, which is more than the population of the provinces of Ontario and Quebec combined.
Each team is named for a common and often seen species seen in our aquatic ecosystems. This blog chapter is about the students and their learning. Surely, you miss seeing some great nature shots, so here is my photo of the Black-shouldered Spinyleg taken at the QUBS dock.
Climate Change Scenarios, Predicted Impacts on Aquatic Ecosystems and Strategies to Mitigate continued Change by Team Cherry-faced Meadowhawks, Sympetrum internum,
Here are some highlights from this team’s presentation.
In the worst case scenario, global average temperature will increase by 4.8 degrees Celsius by 2100. Ice sheets and glaciers will melt, changing northern climates and causing a significant rise in global sea levels. Our carbon dioxide levels will be 3 to 4 times pre-industrial levels. They are already at 400 PPM as measured by NOAA (USA’s National Oceanic and Atmospheric Administration). How do we know that temperature will increase this much? Paleoclimatology provides the answers, principally from ice core analysis. You can see the data well presented at NOAA, where a chart shows over the last 400,000 years natural CO2 changes and temperature changes. Note that the highest measured concentrations were about 300 PPM over this very long period of time.
Our lake ecosystems will be affected greatly, as ice cover in the Great Lakes drops by 42%, meaning species adapted to ice cover and certain temperature ranges will be pressured. River ice will be similarly affected, affecting species diversity, species range and populations. Temperature increases cause more evaporation, increasing salinity and concentrations of other minerals. Oxygen levels will decrease. Carbon dioxide levels will increase, as CO2 is very soluble in water.
Oceans are also absorbing unprecedented concentrations of CO2, increasing ocean acidity. Sea levels have risen an average of 20 centimetres and may rise an additional 1 metre by 2100.
Here, Will tells us about the current effects of climate change. You can read his full list by enlarging the photo by clicking on it.
Many regions in the world will be affected greatly if these changes come to be. The young presenters will feel the full brunt of change and are very concerned. Just a few months ago, China and the USA signed a ground-breaking protocol on climate change. China is spending $50 billion on alternative energy sources, replacing its coal-fired generators at a very fast rate. Two very vulnerable regions in China are the water-scarce North China Plain, and the flood-prone Poyang lake Region.
This graphic shows simply how ice melt will result in a vicious circle of increased heating and further loss of ice.
Fan explains some of the ways we may be able to mitigate and adapt to climate change.
The Intergovernmental Panel on Climate Change (IPCC) had produced a book-sized report in 2012 on this topic.
I found it appropriate somehow that the seminar room was made uncomfortable by the heat and humidity outside. The facts about climate change are also uncomfortable. Perhaps you have heard Australia’s Will Flannery talking about climate change. He is Chief Councillor of Australia’s Climate Council. It started as a government organization and is now a non-governmental organization financed by donors. If you do want more than a summary, then go to its website.
If you wish to delve into the details of the science around climate change, you may refer to the IPCC web site.
QUBS hosts a long-term climate monitoring project (since 2009) using web cameras which monitor forest change. The project started with 12 cameras, 2 in Ontario, 1 at Qubs, and 10 more in the northeastern USA.
Biological Indicator Species by Team Dryad’s Saddle
There are four kinds of indicator species:
Keystone species which create their own ecosystem. Our Beaver (Castor canadensis) comes to mind. One student suggested that the Anchovy, an estuarine brackish water fish common in Chinese waters, is a Chinese candidate.
Flagship species which act as a social target for conservation action. The Chinese have the Panda. We have the Loon.
Sentinel species are the “canary in the coal mine”. Changes to these species tell us there is an environmental concern. Southern species spreading into our latitudes may qualify. Some dragonflies, birds and butterflies previously unheard of in our region are examples, such as Easter Amberwing dragonfly (Perithemis teneris), Red-bellied Woodpecker (Melanerpes carolinus), and the Giant Swallowtail butterlfy (Papilio cresphontes).
Umbrella species protect other species through their activities. The beaver qualifies again. Some African/South American termites also qualify as their giant mounds bring nutrients up from the depths, creating significant areas where other species may thrive. Professor Lougheed has written a paper for this blog on these industrious social insects.
QUBS presents a series of seminars through the summer. They are most informative and entertaining. This year’s series ended Wednesday, August 26th with Stephen Lougheed’s favourite species. Some of his early research projects took Steve to southern South America, where he learned to appreciate the Rufous-Collared Sparrow, Zonotrichia capensis. I cannot possibly do justice to Steve’s love of this species in this blog. His enthusiasm has generated motivation to experience these birds personally. For now, I suggest we experience vicariously through the Web: Rufous Collared Sparrow
What follows is another good example of one of these seminars on July 29, 2015.
James Sinclair of Queen’s University:
James’ topic: Strength in Size or Numbers – disentangling the factors involved in the establishment of non-native species
After dinner in the cafeteria in the Raleigh Robertson Biodiversity Centre, the class, local community members and I convened in the seminar hall in the basement. The hall is set up to accommodate about 75 people, and it was almost full.
James, in quest of an advanced degree, is studying a most pressing topic in ecology: invasive species. Since World War 2, the geometric scale human population increase, combined with ever expanding trade and human migration, has afforded many European and Asian species the opportunity to expand into the Americas (and vice versa!). Most of us have heard about exotic species getting a foothold before this era. The European Starling (Sturnis vulgaris) comes to mind. This feisty, intelligent bird was introduced in the 1890s in New York City by the American Acclimatization Society in its quest to have every bird mentioned in Shakespeare’s works among us in North America (Starlings are mentioned in Henry IV part 1). These same enterprising souls also gave us the House Sparrow (Passer domesticus).
Scientific American gave us on article on the Starling’s origin in North America.
James’ invasive species research focuses on one of our most recent cargo ship hitchhikers, the bloody-red mysid (Hemimysis anomala) which is a shrimp-like crustacean in the Mysida order, native to the Ponto-Caspian region, which has been spreading across Europe since the 1950s and is now in our own St. Lawrence River.
The theoretical basis for James’ research is the concept of propagule pressure. A propagule is a vegetative structure (bud, stem) from a plant from which new plants of the same species will spread. Therefore it is a way to propagate a species. The Red Mangrove (Rhizophora mangle) has populated the shores of the tropics and subtropics in this fashion.
Propagule pressure is a measure of the numbers of a species introduced into a region where they are not native. Since this is a composite measure, you have to know how many were introduced each time, and then how many introductions occurred and over what area. As you can see from the slides taken from James’ presentation below, this is stated as “The set of individuals introduced” and “The rate of introductions”.
There is a minimum population required to “launch” a species and each is different. Only scientific research which emanates from these theoretical underpinnings will clarify how species get started, and this could help us develop more effective strategies for prevention and/or elimination of unwanted species.
As you can see from James’ summary, we have learned a little and have a long way to go to better understand the invasion process.
James’ research target, the small crustacean which you can see in the slide below, has already managed to make the St. Lawrence River its home. The port of Montreal seems to be its “drop-off” point, so the intrepid James decided to sample the port waters to collect research subjects. The best time to collect these light-sensitive crustaceans is at night, in a most seedy part of Montreal’s harbour front. Putting his life on the line, like so many courageous biologists, James was successful in bringing back sufficient mysids to conduct his experiments in the tanks at Queen’s.
For a full listing of events at the Station, including the Wednesday evening summer seminar series, click here.
In Chapter 3 we will experience another dollop of ecological learning from Professor Yuxiang Wang, and see some of the field trips which the students experienced. Some of the facilities at QUBS will be featured. This will be the final chapter for this field course. Tree Swallows research is the next topic, and look for stories and photos featuring Professor Emeritus Raleigh Robertson, a long-time Director of QUBS who started Tree Swallow research at Queen’s in the 1970s.
Unless otherwise credited, all photos are taken by Art Goldsmith.
There is little in life more energizing than being amongst great young minds exploring, studying and testing some of the more pressing questions of today’s world. Such was my opportunity when Queen’s University biology professors Stephen Lougheed and Yuxiang Wang invited me to be with them as they led the Canadian version of the following course at the Queen’s University Biological Station.
Effects of Human Development on Aquatic Environments and Biodiversity in Canada and China Field Course 2015
Aided by research associate Mark Szenteczki, Queen’s grad students Mingzhi Qu and Wenxi Feng, Lougheed and Wang provide Chinese and Canadian undergraduate biology and environmental science students with an opportunity for intensive learning in the field. This learning is mixed with a joyful and exhausting itinerary through some of our country’s large and heavily populated aquatic systems. Learning continues into the evenings with seminars and lectures by course leaders, other biologists and ecologists, and by the students themselves, working in teams.
Partnering with China’s Tongji University, Queen’s University has developed the Sino–Canada Centre for Environment and Sustainable Development, with the Biological Station being the Canadian portion of the Centre.
Although it predates establishment of the Centre, the course, which began in 2005, reflects the Centre’s spirit and its goals. The course is given in summers alternating yearly between China and Canada.
While I experienced only several days of the two-week course at the Biological Station, thanks to material provided by professors Lougheed and Wang, Teaching Assistant Szenteczki and the students, the following includes personal observations, as well as events outside those days when I was present. I have divided my observations into several chapters. In no way is this information comprehensive. Rather, my intent is to provide you, dear reader, with an overview that skims the surface of the wealth of detailed knowledge packed into this very richly composed course.
Course Day 1
The field course provides a rich diversity of experience. On Day 1 at QUBS, the students enjoyed learning about Eastern Ontario natural history and avian diversity. They ended the day with a nocturnal field trip around the Station where they experienced owls, frogs and the numerous insect species which emerge after dark. This blog isn’t intended to give a full annotated itinerary of the course, but rather provide some flavours and snippets of course experiences and content.
First up, a hike at the Station on the Cow Island Marsh Trail.
Any aquatic ecology course has to consider the most productive biological systems—wetlands.
Classification of wetlands is, itself, an interesting and diverse field of study. For the purpose of this blog, we will focus on four classes: marshes (fresh and saltwater, and those in between); fens; bogs; and swamps. Wetland definitions are tenuous and these common names differ from place to place. Much like common bird or plant names, the terms change.
Marshes occur in and along ponds, lakes and rivers; in fact, they occur in and along many aquatic environments. Defined by rich natural nutrient sources, herbaceous emergent vegetation and a neutral pH, marshes are highly ecologically productive places with a diversity of plant and animal life. Of course, they are usually wet! That is, the soils of marshes are usually saturated and overlain by water. There are tidal and non-tidal marshes. Marine marshes are a particular favourite of mine. More about that later.
The Cow Island Marsh is an excellent example. Like so many local marshes in Eastern Ontario, this one is dominated by Common Cattails, Typha latifolia, seen below.
Look closely, though. Increasingly, I have noticed another similar species, Narrow-leaved Cattails, Typha angustifolia, becoming more common and even dominant in some marshes.
On July 29, 2015, Instructor Dale Kristensen of Queen’s University led a walk at Cow Island Marsh that focused on his theme: Plant diversity, identification & importance.
Thanks to the members of team “Scrambled-egg slime mold,” Fei Jin (Fudan University), Zixiang Li (Beijing Normal University), Sarah Minnes (Memorial University) and Natalie Wong (University of Toronto), for their write-up on this event. Thanks go out to Mark Szenteczki as well for the two photos showing Dale leading the students at Cow Island Marsh.
Some of the plants and scenes observed at this marsh.
Swamp Milkweed, Asclepias incarnata, which is not as familiar as its field growing cousin, Common Milkweed. It is, though, a favourite also of many butterflies.
I had not noticed the Marsh Bellflower, Campanula aparinoides, before. Dr. Kristensen identified it immediately as a common local marsh inhabitant. It is sometimes overlooked because of its diminutive size and vine growth habit that often causes the majority of the long narrow leaves to be hidden by other plants
To the right are the stem and leaves of the same plant stretched across the boardwalk to enable photography.
Note that the plant was not harmed during this process!
Many odonates (dragonflies and damselflies) inhabit the marsh in midsummer.
One common dragonfly is the Twelve-spotted Skimmer, Libellula pulchella (photo below).
At the entrance to the boardwalk, in July, you may see a most symmetrical flower, the Buttonbush, Cephalanthus occidentalis, a wetland-loving member of the Madder family.
Invasive species, a subject of a talk given by James Sinclair at QUBS during the China–Canada course, are apparent in the marsh. Look for another posting featuring Sinclair’s presentation. Though the Purple Loose-strife (Lythrum salicaria) is now controlled, the plant below, the European Frog-bit, Hydrocharis morsus-ranae, is invading most of our marshes.
About 23 years ago, the Ontario Ministry of Natural Resources and Forestry (OMNRF), in partnership with the Ontario Federation of Anglers and Hunters (OFAH), established an Invading Species Awareness Program, where you can learn more about this species and how to control it, as well as the growing number of species invading Ontario.
The lovely, and invasive, Flowering Rush, Butomus umbellatus, Westmeath Provincial Park, Ontario.
Human effects on all wetlands have reduced these important ecosystems both qualitatively and quantitatively. This photo permits us a more sanguine view, perhaps echoing a previous time when the marsh and its human inhabitants lived more harmoniously. The marsh is in the foreground, Cow Island on the upper left and Lake Opinicon beyond.
The course focuses on freshwater systems. The Queen’s University Biological Station includes properties in the Frontenac Axis, a band of the Canadian Shield that extends from the Algonquin Highlands across the St. Lawrence River into New York State, where the band widens to form the Adirondacks. Swamps, bogs, marshes and fens are a feature of the rocky forested landscape. Locally, swamps and marshes are well represented. One of the best fens in the area is the White Lake Fen, near Arnprior, Ontario.
Fens receive groundwater, and, therefore, are more nutrient rich and biodiverse than bogs which receive only rainwater. Both are characterized by both herbaceous and woody water-loving plants, including many orchid and carnivorous plant species.
Swamps are characterized by woody vegetation. Cedar swamps abound in Eastern Ontario. Eastern White Cedars, Thuja occidentalis, tend to be some of the oldest trees in our country. Drainage has left a great deal of our cedar swamps with a lowered water table, which has caused a drop in diversity and no cedar regeneration. Cedars are very adaptable, though, and upland populations are increasing as they invade abandoned farmlands. This points out a problem with the way we organize our conservation efforts around endangered species instead of endangered ecosystems. The cedar is definitely not endangered. Perhaps the cedar swamp is threatened?
Pictured on the right is a cedar grove in Stittsville, Ontario. Previously, this grove, now a protected area, had standing water most of the year.
Bogs and fens are indeed a northern phenomenon. In Eastern Ontario, well known large bogs exist and even have moose populations (Alfred Bog and Mer Bleue). Just for fun, and because your blogger recently completed a lifelong dream trip to a very southern bog, here is a photo from the Okefenokee National Wildlife Refuge (southern Georgia, USA). Bogs’ waters are only replenished by rain. This makes them nutrient poor, acidic, wet environments characterized by peat moss. The southern climate produces some bigger trees, and some more diversity than one would get in our local bogs. Still, Okefenokee is NOT a swamp.
Course Day 2
Each team wrote their own blog about each day of the course. For the Day 2 content, thanks go out to team ‘Dryad’s saddle’ members, Derek James Newton (Queen’s), Qin Lanxue (Tongji), Xing Kangnan (BNU) and Lyu Wenyang (d’Overbroecks).
Before breakfast, the group hiked, working up an excellent appetite, looking for some of the many species of birds resident in the marshes, forests, lakes and shores surrounding QUBS. Along the way, they learned a little about the Grey Rat Snake, Pantherophis spiloides, our largest snake in Ontario and endemic to the Frontenac Axis. This threatened reptile is often seen moving through the property.
Indeed, this blogger encountered the snake below on the same road the students hiked (right).
The students heard a Pine Warbler, Setophaga pinus; many black-capped chickadees, Poecile atricapillus; and they heard the sharp “chick-chick” calls from a Downy Woodpecker, Dryobates pubescens. As they left the forest on their way to the marsh, they observed Common Yellowthroat warblers, Setophaga dominica; and Blue Jays, Cyanocitta cristata. Walking along the marsh boardwalk, the students heard the “prehistoric caw” (note: the blogger thinks of this loud abrupt call as a “groink”) of the Great Blue Heron, Ardea herodias. They saw a Caspian tern, Hydroprogne caspia, fly over as it fished Lake Opinicon. The first true wetland resident species encountered was the Swamp Sparrow, Melospiza georgiana, which popped up in the bullrushes and cattails. Other species usually heard or seen around the lake are the Common Loon, Gavia immer; and the Osprey, Pandion haliaetus.
During the afternoon, the group learned about Global Positioning Systems (GPS) and applications to environmental science research. Over the last 20 years, GPS and Geographic Information Systems, in combination with remote sensing, have become fundamental tools for learning about, conducting research on and presenting clear visualizations of environmental topics.
Qu Mingzhi provided comprehensive knowledge on GPS methods and applications.
Following a walk to an upland marsh, dotted with willow and goldenrod, the students were treated to another presentation by a Queen’s grad student Wenxi Feng, who is working on the applications of monitoring for eDNA. Wenxi also presented his research at the QUBS Open House in June, which your blogger attended. The idea is simple; the application is much more complex. Wetland organisms, such as fish, turtles and frogs, for example, through normal life processes, exude mitochondrial DNA. Water samples may be analyzed for this DNA indicating presence or absence, density, and much more information about organisms in the ecosystem, without the need to capture or harvest the organisms.
Wenxi Feng showed course participants eDNA methods and applications for environmental research. This is a developing and exciting field, which has great potential for streamlining and improving environmental monitoring.
The day ended with participants watching one of my favourite motion pictures, The Big Year. Three American “birders” compete to see the most bird species in a single year. Of course they are all men, who go to great lengths to find that rare bird.
With that, this first chapter of the 2015 China–Canada Field Course ends. In the next chapter, we will follow the participants as they develop their own seminars and I will give details about several of the student seminars.
Thanks to Janice Tripp for her expert editing assistance.
Professor Stephen Lougheed started this blog in 2009.
It is intended to be a vibrant and factual resource for learning and documenting the science and natural history of the Queen’s University Biological Station (QUBS) and surroundings. As you can see from recent postings, the content also ranges geographically since people who have frequented QUBS now span the Earth. People working at QUBS carry on research in many other lands and ecosystems.
My plan is to add a further dimension to an already successful venture: the people, facilities and the various happenings at the Station. There is a dynamic to the natural history and science that becomes more real, informative and lasting if we also know more about the people generating all that good science and knowledge. So, look for more postings about the growing buffet of field courses and ongoing research projects.
Submissions are welcome, subject to editorial assessment – send to Stephen Lougheed.
Stephen is always around and involved. His busy schedule as a Full Professor at Queen’s University and Director at the Station doesn’t leave sufficient time for him to always capture all of what is going on at QUBS. Therefore, I volunteered to take a lead role in producing the blog content.
Allow me to introduce myself. Most of my spare time is dedicated to natural history and environmental knowledge development and communication.
I hold an executive position with the Macnamara Field Naturalists’ Club of Arnprior, Ontario, which is where I live. We have the tallest tree in Ontario there, in Gillies Grove. It is a White Pine (Pinus strobus) with a height of about 47 metres. The National Research Council of Canada has placed me on their Animal Care Committee, which oversees the treatment and care of laboratory animals in human health research.
I have my own blog, which captures natural history knowledge and more as I make my way through various eco-districts.
It is best to READ the blog rather than describing it, as it will also give you the flavour of future Opinicon Natural History postings.
I worked at Environment Canada from 1980 to 1997, as Chief and then Director of Conservation Service Policy and Knowledge Integration, where some of my achievements included the development of a large portion of Canada’s Green Plan and Environment Canada’s entry onto the World Wide Web.
Between 1980 and 1988, I developed and advised on water policy and legislation, managed Lower Fort Garry National Historic Park and led social development policy for the Government of Manitoba.
All said and done, I have returned to my first passion—natural history and environmental science. Expect diversity and variety with an emphasis on “interesting.” Comments are welcome. Please point out errors and omissions. Most of all, keep returning and let me know what you think.
In the next few weeks, look for editions about:
1. Cow Marsh Nature Trail
2. What’s Up with the Tree Swallow Boxes at QUBS?
3. Sino-Canada Eco-dreams OR Effects Of Human Development On Aquatic Environments and Biodiversity In Canada And China
4. The Bug Man Cometh (Field Entomology & Ecology)
5. Fungi with a Fun Guy (Fabulous Fall Fungi)
6. And your comments, recommendations and contributions!
By S.C. Lougheed
For Ontarions, the word “termite” conjures up a negative image of ravenous insects that cause immense and costly damage to human-made wooden structures because of their propensity to eat dead wood and indeed any material that is cellulose-based (Evans 2011). The beast that we know in Ontario is the eastern subterranean termite (Reticulitermes flavipes), a species native to the eastern USA that has been introduced multiple times into Ontario (Scaduto et al. 2012) – probably first in 1938 (Urquhart 1953).
In other parts of the world, like the savannahs of African savannahs, the pampas of Argentina, or tropical and subtropical Australia, some termite species present another face – that of exquisite natural engineers who create magnificent and sometimes immense structures of cellulose, mud and saliva (Figure 1). These termite mounds afford many benefits to the termite colony including protection from predators and buffering from sometimes extreme environments where they are found. In the Box below Ipresent some basic information on evolutionary affinities and diversity.
One of my favourite examples of beautifully-adapted insect architecture is the mound of the magnetic termite, Amitermes meridionalis, found in Northern Australia. Magnetic termites build their wedge-shaped mounds on seasonal flood plains that are saturated during the wet season (precluding subterranean abodes) and baked in the intense tropical sun in the dry season – an extreme environment indeed! The photo in Figure 2 shows that the mounds are all oriented in the same direction – north-south. The unique shape and orientation mean that one side is shaded and cool as the sun rises and sets, but also that when the sun is at its zenith, only the very top of the wedge receives direct sunlight. Termite mounds can be incredibly important to other organisms. Hollows within them can provide shelter for animals like goannas (monitor lizards), quolls (small marsupials), and snakes. For some species termites form a significant part of their diet (e.g. bilbies – small arid-land omnivorous marsupial) and termite mounds thus a rich foraging ground. Finally termite mounds play a significant role in enriching and cycling of nutrients, with local effects persisting decades after a colony has disappeared.
Evans, T.A. 2011. Invasive termites, pp. 519-562. In D.E. Bignell, Y. Roisin, & N. Lo Eds., Biology of Termites: A Modern Synthesis. Springer, Dordrecht, the Netherlands.
Scaduto D.A., S.R. Garner, E.L. Leach & G.J. Thompson. 2012. Genetic evidence for multiple invasions of the eastern subterranean termite into Canada. Environ. Entomol. 41: 1680-168.
Urquhart, F.A. 1953. The introduction of the termite into Ontario. Can. Entomol. 85: 292-293.
Box. There are over 3000 named species of termites (also called “white-ants”), although undoubtedly there remain many others to be discovered (Krishna et al. 2013). Much of this species richness is centred in the tropics and subtropics, where termites play a major role in ecosystems as detritivores. Originally placed within their own order (Isoptera), recent molecular evidence suggests that termites are most closely allied to cockroaches with suggestions that Isoptera be subsumed within the cockroach order Blattodea (Inward et al. 2007). Termites are eusocial insects where different castes perform different roles within the colony. This phenomenon of eusociality has arisen multiple times both in insects (e.g. Hymenoptera – bees and wasps), in crustaceans (alpheid snapping shrimp), and in mammals (naked mole rats, Heterocephalus glaber).
Inward, D., G. Beccaloni & P. Eggleton. 2007. Death of an order: A comprehensive molecular phylogenetic study confirms that termites are eusocial cockroaches. Biol. Lett. 3: 331-335.
Krishna, K, D.A. Grimaldi, V. Krishna & M.S. Engel. 2013. Treatise on the Isoptera of the world. Bull. Am. Mus. Nat. Hist. 377: 1–2704.
The parrots comprise a large order (Psittaciformes) of birds with a mainly pantropical distribution, although some species do inhabit temperate regions in the Southern Hemisphere as well (e.g. the burrowing parrot, Cyanoliseus patagonus, of southern South America). Number of species reported varies but generally is on the order of 340 to 370 distributed across between 78 and 86 genera (Rowley and Collar 1997). Characteristics of parrots will be familiar to most: robust, curved bill, strong legs with zygodactylous feet (two toes forward, two toes facing backward). Many are brightly-coloured although some, like the sulfur-crested cockatoo (Cacatua galerita) are mostly white, while others, like the flightless Kakapo (Strigops habroptila) of New Zealand, have muted and cryptic plumage patterns to avoid predators.
In my still unfolding peregrinations in Australia I have already seen 10 species of parrot including this lovely northern rosella (photo by Cam Hudson – see his blog – from some distance – but still showing some of the vibrant colours).
My list thus far includes:
Red tailed black-cockatoo, Calyptorhynchus banksii
Unfortunately, at least 80 species of parrot are classified as vulnerable or endangered (IUCN 2013) due to a mixture of habitat loss, collection for the pet trade, and persecution because some are considered agricultural pests (Collar 2007) with some already extinct. Indeed, the only psittacid of Eastern North America went extinct in the early 20th Century. The Carolina parakeet (Cacatua galerita) once ranged from southern New York, south to the Gulf of Mexico and as far west as Nebraska (Snyder 2004). The Carolina parakeet was a lovely species, with bright yellow head, orange face, green body and pale bill (see John James Audubon’s rendering here). One can imagine that, before European settlement (and ensuing loss of the Eastern deciduous forest, persecution because it foraged on orchards, and hunting for the millinery trade – nothing like a stuffed parakeet on your hat I guess – see Saikku, 1990), very occasionally one might even have seen a northern vagrant parakeet in Canada.
Collar, N.J. 2007. Globally threatened parrots: criteria, characteristics and cures. International Zoo Yearbook 37: 21–35.
IUCN 2013. IUCN Red List of Threatened Species. Version 2013.2. <www.iucnredlist.org>. Downloaded on 22 January 2014.
Rowley, I. and N.J. Collar. 1997. Order PSITTACIFORMES. In Handbook of the Birds of the World – Volume 4. Sandgrouse to Cuckoos. (J. del Hoyo, A. Elliott, J. Sargatal eds.) Lynx Edicions
Saikku, M. 1990. The extinction of the Carolina parakeet. Environmental History Review 14: 1-18.
Snyder, N.F.R. 2004. The Carolina Parakeet: Glimpses of a Vanished Bird Princeton University Press. Princeton, NJ.