References for Commentary on FoxP2 in Bats:
1. Li G, Wang J, Rossiter SJ, Jones G, Zhang S (2007) Accelerated FoxP2
Evolution in Echolocating Bats. PLoS ONE 2(9): e900.
doi:10.1371/journal.pone.0000900
http://www.plosone.org/article/info:doi%2F10.1371%2Fjournal.pone.0000900
2. Pettigrew, J.D. and Kirsch, J.A.W. (1998) The bat problem. I.
DNA-hybridisation melting curves based on DNA enriched for AT- or
GC-content. Phil. Trans. Roy. Soc. B, 353: 369-379.
3. Kirsch, J. A. W. & Pettigrew, J. D. (1998) Base
compositional biases and the bat problem. II.
DNA-hybridisation trees based on tracers enriched for AT- or GC-
content. Phil. Trans. Roy. Soc. B, 353: 381-388
These two papers use a technique suggested by Roy Britten to melt
DNA at different temperatures so that fractions could be obtained at a
range of temperatures. High temperatures yielded GC-rich DNA,
since G &C are joined by 3 hydrogen bonds and so require higher
temperatures to dissociate or melt, while lower temperatures yielded
AT-rich DNA, since A&T are joined by only two hydrogen bonds and
melt at a lower temperature. Such fractions showed that there was
a large error in estimating the rhinolophoid-megabat distance because
these two groups of bats share the highest AT content in vertebrate
DNA. This distance increased when GC-rich DNA was used, showing that
uncorrected measurements of that distance were under-estimated by a
large margin that would lead to an artifactual association between
rhinolophoids and megabats.
4. Hutcheon, J.M., Kirsch, J.A.W., & Pettigrew, J.D. (1998)
Base-compositional biases and the bat problem. III. the question
of microchiropteran monophyly Phil. Trans. Roy. Soc. B,
353:607-617
This paper is often quoted in support of microbat paraphyly, even
though it completely ignores the evidence from [3 &4] that the
rhinolophoid-megabat affinity is an artifact of the high AT content of
these two otherwise dissimilar groups of bats. This study made no
effort to correct for this artifact during data collection. The late
John Kirsch mischievously set this paper as a trap, to see who would be
influenced more by the tree with pictured taxa in this paper than by
the hard evidence for a biasing effect of high AT content from the
previous two papers by the same lab.
5. Teeling EC, Springer MS, Madsen O, Bates P, O'brien SJ, Murphy WJ.
2005
A molecular phylogeny for bats illuminates biogeography and the
fossil record. Science. 2005 Jan 28;307(5709):580-4.
6. Jones G, Teeling EC.
The evolution of echolocation in bats. Trends Ecol Evol. 2006
Mar;21(3):149-56
7. Teeling EC, Madsen O, Van den Bussche RA, de Jong WW, Stanhope MJ,
Springer MS.
Microbat paraphyly and the convergent evolution of a key
innovation in Old World rhinolophoid microbats. Proc Natl Acad Sci U S
A. 2002 Feb 5;99(3):1431-6.
Papers 5, 6 & 7 put forward a radical new phylogeny of bats,
paraphyly, that is unsupported by virtually every other
discipline apart from DNA sequence data. The association of
megabats and rhinolophoid microbats is put forward as fact despite the
absence of any supporting evidence from other disciplines and despite
the fact that the association is contradicted by molecular evidence
from proteins such as serum proteins [8], hemoglobin,alpha-crystallin,
etc. It would not be possible to find any bats that are more
different than megabats and rhinolophoids, so it is not surprising that
morphological studies emphatically support the molecular protein data
in contradicting paraphyly (e.g. Jones KE, Purvis A, MacLarnon A,
Bininda-Emonds, Simmons NB.(2002) A phylogenetic supertree of the
bats (Mammalia: Chiroptera).Biol Rev Camb Philos Soc. 77(2):223-59).
The source of the error is not hard to find, since megabats and
rhinolophoids have the most modified genomes of any vertebrate. Seen in
the light of the isochore phenomenon, these two groups of bats will
have extremely similar DNA sequences in many genes that are not
otherwise protected in the H3 isochore, like FoxP2. Convergent
similarity of DNA sequences is a real phenomenon, despite derisory
comments to the contrary, and is an unpleasant feature that requires
uusual measures to overcome it, such as the use of translated protein
sequence data or innovative distance techniques. The cursory
measurements of overall base composition that were used in
studies 5 & 7 would not be adequate to uncover the site -specific
changes in DNA base composition underlying isochore formation.
8. Schreiber A Bauer D and Bauer K 1994 Mammalian evolution from serum
protein epitopes Biological Journal of the Linnaean
Society51:359-376
This is an example of the many studies using proteins that are quite
incompatible with the new paraphyly of microbats that links
rhinolophoids to megabats. While not all such studies explicitly
address the bat problem, it is obvious that protein sequence data from
hemoglobin, alpha-crystallin, opsins, etc fail to make this anomalous
linkage between the two most contrasting of any bat groups. One wonders
why the protein is ignored in favour of the DNA, which is strongly
biased in these two groups of bats. It is worth noting that conversion
to amino acid codons can overcome DNA biases, a correction not applied
in any of the DNA sequence studies that claim a paraphyletic
arrangement between rhinolophoid microbats and megabats.
9. Frances E. Arrighi1, William Z. Lidicker Jr.2, Manley Mandel1 and
Janet Bergendahl1Biochemical Genetics 6: 27-30 Heterogeneity in CsCl
buoyant densities of chiropteran DNA
10. Sabeur G, Macaya G, Kadi F, Bernardi G The isochore
patterns of mammalian genomes and their phylogenetic implications. J
Mol Evol. 1993 Aug;37(2):93-108
<>11. Naylor GJP, Brown WM SYSTEMATIC BIOLOGY 47: (1) 61-76
MAR 1998
Amphioxus mitochondrial DNA, chordate phylogeny, and the limits of
inference based on comparisons of sequences
12. Loomis WF, Smith DW. 1995 Experientia.51: 1110-1115 Consensus
phylogeny of Dictyostelium.
<>
14. Yager, David D. (1989) Audition in the praying mantis,Mantis
religiosa L.: identification of an interneuron mediating ultrasonic
hearing. Journal of Comparative Physiology A 165(4)
<> 15. L F. GÁLL LF1 and. TIFFNEY BF 1983 Science
219: 507-509 A Fossil Noctuid Moth Egg from the Late Cretaceous of
Eastern North America
<> 16. Pierson, ED. 1986 Molecular Systematics of the
Microchiroptera: Higher taxon relations and biogeography. PhD Thesis.
UC Berkeley.
<> 17. Summary of FoxP2 substitutions in bats.
2
Pteropus
V L
7 Cynopterus I V M H G
G I
6 Rhinolophus A P A S A T
6
Asellicus I
D P M S T
3 Hipposideros I T M
9
Coelops
V I D M D T A L S
13 Megaderma N T S S A A P A T V N T S
3 Taphozous L G L
8
Nycteris A
T A A F E V S
3 Pteronotus I M T
4
Carollia
M L T S
12 Vesper bats M V I V N L N A V L P M
1 Chaerophon V
3
Whales
P A M
If microbats diverged 100 MyA and whales ~40 MyA,
then there should be 7-8 substitutions in
microbats. Actual value is:-MEAN = 5.7 (range 1-13). The
prediction
would be more accurate if it were known whether
all the whales inherited their complement of
FoxP2 changes from an ancestor (as seems likely, since both
echolocating odontocetes and non-echolocating mystecetes have the same
three FoxP2 substitutions). The older age of this ancestor would align
the prediction. Note that these calculations would make some microbats
very old indeed, such as the vespertilionids, rhinolophoids,
megadermatids and nycterids. No wonder there is so much strife in
elucidating a phylogeny for some microbats!