To: Meril, Rick
Sent: Wed May 25 12:13:46 2011
Subject: RE: Fwd: FW: new canid genetics paper
Hi Rick,
Tyler, Linda & I first heard Roland present these results at the Midwest wolf Stewards meeting in Wisconsin about a month ago and were certainly both interested and intrigued. While we remain open to rigorous testing of the ideas suggested by this work re the origins and taxonomy of eastern wolves (C. lycaon), we are not yet convinced and have concerns that impact the conclusions of vonHoldt et al. concerning the eastern wolf.
One of our primary concerns is that the authors refer to eastern wolves and Great Lakes wolves interchangeably, and conduct their subsequent analyses based on this assumption. We must assume some of the blame for this confusion given suggestions in earlier works from this lab of a broad range for eastern wolves (e.g. Grewal et al. 2004; J. Mammalogy 85: 625-632), but in recent papers (e.g. Rutledge et al. 2010: Heredity 105: 520-531; Wheeldon et al. 2010: Molecular Ecology 19: 4428-4440) we tried to clarify that: 1) while there are likely no remaining unhybridized eastern wolves in the wild, the closest living relatives to the historic eastern wolf live in and immediately around Algonquin park in central Ontario. These wolves are distinct from wolves in NE and NW Ontario, as well the Great Lakes States, and 2) Although wolves in the Great Lakes States and much of Ontario and Quebec contain some eastern wolf genetic material, they are not eastern wolves and both phenotypically and genetically (based on autosomal microsatellites) group more closely with Gray wolves, C. lupus than with C. lycaon (again, as typified by wolves in Algonquin). Given this, the finding that Great Lakes wolves and Red wolves did not share a common evolutionary origin is not surprising. We have suggested a common origin for Red wolves and Eastern wolves (again typified by, and largely restricted to wolves in Algonquin), NOT between Red wolves and the Great Lakes wolf which, as mentioned above, is a hybrid of C. lupus and C. lycaon.
Given that VonHoldt et al. only analyzed DNA from 2 Algonquin wolves, and that we don't know when and exactly where these samples were collected (i.e. they could have been coyotes or hybrids collected somewhere in or around Algonquin), we don't believe that the hypothesis of eastern wolves as a distinct North American evolved species was adequately assessed by this work. Note also that the Wilson et al. (2000) canid evolutionary model (CJZ 78: 2156-2166) of the eastern wolf suggests divergence from the western coyote only 150-300K years ago. This time is barely sufficient to see differences in the mtDNA control region resulting from mutation, so the finding that genomic SNPs did not differentiate eastern wolves from western coyotes is not surprising. Another concern relates to the analyses conducted using the program Structure. Anyone familiar with this program, used to assign membership to different genetic groups, knows that it would be highly unlikely for any "population" consisting of only 2 individuals to separate as a distinct group from other larger populations. Nonetheless, it is interesting that the PCA conducted by VonHoldt et al. (see their Fig . 3) places the 2 Algonquin samples separate from all other groups although the authors arbitrarily grouped them with Great Lakes wolves.
Future research might yet reveal that there never was a North American evolved Red wolf or Eastern wolf, and that these animals are indeed merely hybrids between C. lupus and C. latrans, but if so we wonder how the following lines of evidence supporting a North American evolved wolf distinct from the Gray wolf will be rectified:
1) Hybridization between eastern wolves/ red wolves and coyotes is pervasive where they are sympatric in eastern North America but hybridization between wolves and coyotes remains exceedingly rare or absent in the west. Hybridization between wolves and coyotes is also very rare across the Western Great Lakes region (see Wheeldon et al. paper cited above), and in northern Ontario (east and west). The range of ratios of abundance of wolves: coyotes vary widely in both eastern and western North America so saying the 2 species only hybridized in the east because of skewed species ratios requires quite a leap of faith.
2) How does one explain the presence of mitochondrial haplotypes C3 and C13 (see Rutledge and Wheeldon refs cited above); both of which are common in eastern wolves and their associated hybrids, but neither of which are found in non-hybridizing wolves or coyotes (i.e. gray wolves and western coyotes).
3) Evidence of a separate Y-chromosome eastern wolf lineage (Wilson et al., manuscript in review).
In summary, while we agree that the approaches employed by VonHoldt et al. represent an important step forward re analysis of canid taxonomy; until a more representative and balanced sample containing eastern wolves (i.e. Algonquin wolves), historic pre-Columbian eastern wolf samples, and the appropriate out groups, is similarly analyzed we consider the hypothesis of a North American evolved wolf independent of the gray wolf still viable.
Cheers,
Brent
Brent Patterson
Research Scientist – wolves and deer
Adjunct Professor, Trent University, Environmental and Life Sciences Graduate Program
President, Ontario Chapter of The Wildlife Society
Ontario Ministry of Natural Resources
Wildlife Research and Development Section
Trent University, DNA Building
2140 East Bank Drive
Peterborough, ON
K9J 7B8, CANADA
Tel: (705) 755-1553
Fax: (705) 755-1559
http://people.trentu.ca/brentpatterson/
Tyler, Linda & I first heard Roland present these results at the Midwest wolf Stewards meeting in Wisconsin about a month ago and were certainly both interested and intrigued. While we remain open to rigorous testing of the ideas suggested by this work re the origins and taxonomy of eastern wolves (C. lycaon), we are not yet convinced and have concerns that impact the conclusions of vonHoldt et al. concerning the eastern wolf.
One of our primary concerns is that the authors refer to eastern wolves and Great Lakes wolves interchangeably, and conduct their subsequent analyses based on this assumption. We must assume some of the blame for this confusion given suggestions in earlier works from this lab of a broad range for eastern wolves (e.g. Grewal et al. 2004; J. Mammalogy 85: 625-632), but in recent papers (e.g. Rutledge et al. 2010: Heredity 105: 520-531; Wheeldon et al. 2010: Molecular Ecology 19: 4428-4440) we tried to clarify that: 1) while there are likely no remaining unhybridized eastern wolves in the wild, the closest living relatives to the historic eastern wolf live in and immediately around Algonquin park in central Ontario. These wolves are distinct from wolves in NE and NW Ontario, as well the Great Lakes States, and 2) Although wolves in the Great Lakes States and much of Ontario and Quebec contain some eastern wolf genetic material, they are not eastern wolves and both phenotypically and genetically (based on autosomal microsatellites) group more closely with Gray wolves, C. lupus than with C. lycaon (again, as typified by wolves in Algonquin). Given this, the finding that Great Lakes wolves and Red wolves did not share a common evolutionary origin is not surprising. We have suggested a common origin for Red wolves and Eastern wolves (again typified by, and largely restricted to wolves in Algonquin), NOT between Red wolves and the Great Lakes wolf which, as mentioned above, is a hybrid of C. lupus and C. lycaon.
Given that VonHoldt et al. only analyzed DNA from 2 Algonquin wolves, and that we don't know when and exactly where these samples were collected (i.e. they could have been coyotes or hybrids collected somewhere in or around Algonquin), we don't believe that the hypothesis of eastern wolves as a distinct North American evolved species was adequately assessed by this work. Note also that the Wilson et al. (2000) canid evolutionary model (CJZ 78: 2156-2166) of the eastern wolf suggests divergence from the western coyote only 150-300K years ago. This time is barely sufficient to see differences in the mtDNA control region resulting from mutation, so the finding that genomic SNPs did not differentiate eastern wolves from western coyotes is not surprising. Another concern relates to the analyses conducted using the program Structure. Anyone familiar with this program, used to assign membership to different genetic groups, knows that it would be highly unlikely for any "population" consisting of only 2 individuals to separate as a distinct group from other larger populations. Nonetheless, it is interesting that the PCA conducted by VonHoldt et al. (see their Fig . 3) places the 2 Algonquin samples separate from all other groups although the authors arbitrarily grouped them with Great Lakes wolves.
Future research might yet reveal that there never was a North American evolved Red wolf or Eastern wolf, and that these animals are indeed merely hybrids between C. lupus and C. latrans, but if so we wonder how the following lines of evidence supporting a North American evolved wolf distinct from the Gray wolf will be rectified:
1) Hybridization between eastern wolves/ red wolves and coyotes is pervasive where they are sympatric in eastern North America but hybridization between wolves and coyotes remains exceedingly rare or absent in the west. Hybridization between wolves and coyotes is also very rare across the Western Great Lakes region (see Wheeldon et al. paper cited above), and in northern Ontario (east and west). The range of ratios of abundance of wolves: coyotes vary widely in both eastern and western North America so saying the 2 species only hybridized in the east because of skewed species ratios requires quite a leap of faith.
2) How does one explain the presence of mitochondrial haplotypes C3 and C13 (see Rutledge and Wheeldon refs cited above); both of which are common in eastern wolves and their associated hybrids, but neither of which are found in non-hybridizing wolves or coyotes (i.e. gray wolves and western coyotes).
3) Evidence of a separate Y-chromosome eastern wolf lineage (Wilson et al., manuscript in review).
In summary, while we agree that the approaches employed by VonHoldt et al. represent an important step forward re analysis of canid taxonomy; until a more representative and balanced sample containing eastern wolves (i.e. Algonquin wolves), historic pre-Columbian eastern wolf samples, and the appropriate out groups, is similarly analyzed we consider the hypothesis of a North American evolved wolf independent of the gray wolf still viable.
Cheers,
Brent
Brent Patterson
Research Scientist – wolves and deer
Adjunct Professor, Trent University, Environmental and Life Sciences Graduate Program
President, Ontario Chapter of The Wildlife Society
Ontario Ministry of Natural Resources
Wildlife Research and Development Section
Trent University, DNA Building
2140 East Bank Drive
Peterborough, ON
K9J 7B8, CANADA
Tel: (705) 755-1553
Fax: (705) 755-1559
http://people.trentu.ca/brentpatterson/
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From: Bradley White <bradley.white@sympatico.ca>
To: Meril, Rick
Sent: Sun May 22 14:27:52 2011
Subject: RE: Fwd: FW: new canid genetics paper
To: Meril, Rick
Sent: Sun May 22 14:27:52 2011
Subject: RE: Fwd: FW: new canid genetics paper
Hi Rick:
We still interpret the data such that the eastern/red wolf are distinct but closely related species to the western coyote. Wayne et al are still making interpretations about coyote /gray wolf hybridization to support the original Wayne suggestion that the red wolf was formed a hybrid.
We still interpret the data such that the eastern/red wolf are distinct but closely related species to the western coyote. Wayne et al are still making interpretations about coyote /gray wolf hybridization to support the original Wayne suggestion that the red wolf was formed a hybrid.
The Biology is clear; coyotes and gray wolves do not hybridise. The gray/eastern hybrids do not hybridize with either eastern or western coyotes. Coy wolves or eastern coyotes have little to no gray wolf material.
Brad
-___________________________________________________________
From: Ron Nowak <ron4nowak@cs.com>
To: Meril, Rick
Sent: Sun May 22 11:24:25 2011
Subject: Re: new canid genetics paper
To: Meril, Rick
Sent: Sun May 22 11:24:25 2011
Subject: Re: new canid genetics paper
Rick---it's just more of the same; the paper is filled with qustionable material and conclusions. And as long as there is another major team of geneticists taking a diametrically opposite view, I cannot accept the validity of that whole approach---Ron
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