PMID- 18198331OWN - NLMSTAT- MEDLINEDA - 20080116DCOM- 20080313LR - 20081121IS - 0890-9369 (Print)VI - 22IP - 2DP - 2008 Jan 15TI - Understanding of bat wing evolution takes flight.PG - 121-4AD - Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.FAU - Cooper, Kimberly LAU - Cooper KLFAU - Tabin, Clifford JAU - Tabin CJLA - engGR - F32 HD 052349/HD/NICHD NIH HHS/United StatesGR - R37 HD 32443/HD/NICHD NIH HHS/United StatesPT - CommentPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPL - United StatesTA - Genes DevJT - Genes & developmentJID - 8711660RN - 0 (Homeodomain Proteins)SB - IMCON - Genes Dev. 2008 Jan 15;22(2):141-51. PMID: 18198333MH - AnimalsMH - Chiroptera/*geneticsMH - *EvolutionMH - Forelimb/anatomy & histologyMH - FossilsMH - *Genetic VariationMH - Homeodomain Proteins/*geneticsMH - Wing/*growth & developmentEDAT- 2008/01/17 09:00MHDA- 2008/03/14 09:00CRDT- 2008/01/17 09:00AID - 22/2/121 [pii]AID - 10.1101/gad.1639108 [doi]PST - ppublishSO - Genes Dev. 2008 Jan 15;22(2):121-4.

PMID- 16618938OWN - NLMSTAT- MEDLINEDA - 20060427DCOM- 20060615LR - 20081120IS - 0027-8424 (Print)VI - 103IP - 17DP - 2006 Apr 25TI - Development of bat flight: morphologic and molecular evolution of bat wing digits.PG - 6581-6

AB - The earliest fossil bats resemble their modern counterparts in possessing greatly elongated digits to support the wing membrane, which is an anatomical hallmark of powered flight. To quantitatively confirm these similarities, we performed a morphometric analysis of wing bones from fossil and modern bats. We found that the lengths of the third, fourth, and fifth digits (the primary supportive elements of the wing) have remained constant relative to body size over the last 50 million years. This absence of transitional forms in the fossil record led us to look elsewhere to understand bat wing evolution. Investigating embryonic development, we found that the digits in bats (Carollia perspicillata) are initially similar in size to those of mice (Mus musculus) but that, subsequently, bat digits greatly lengthen. The developmental timing of the change in wing digit length points to a change in longitudinal cartilage growth, a process that depends on the relative proliferation and differentiation of chondrocytes. We found that bat forelimb digits exhibit relatively high rates of chondrocyte proliferation and differentiation. We show that bone morphogenetic protein 2 (Bmp2) can stimulate cartilage proliferation and differentiation and increase digit length in the bat embryonic forelimb. Also, we show that Bmp2 expression and Bmp signaling are increased in bat forelimb embryonic digits relative to mouse or bat hind limb digits. Together, our results suggest that an up-regulation of the Bmp pathway is one of the major factors in the developmental elongation of bat forelimb digits, and it is potentially a key mechanism in their evolutionary elongation as well.AD - Howard Hughes Medical Institute, Department of Pediatrics, Section of Developmental Biology, University of Colorado at Denver and Health Sciences Center, 12800 East 19th Avenue, Aurora, CO 80045, USA.FAU - Sears, Karen EAU - Sears KEFAU - Behringer, Richard RAU - Behringer RRFAU - Rasweiler, John J 4thAU - Rasweiler JJ 4thFAU - Niswander, Lee AAU - Niswander LALA - engSI - GENBANK/DQ279782SI - GENBANK/DQ279783SI - GENBANK/DQ279784SI - GENBANK/DQ279785GR - F32 HD050042-01/HD/NICHD NIH HHS/United StatesGR - HD32427/HD/NICHD NIH HHS/United StatesPT - Comparative StudyPT - Journal Article

PT - Research Support, N.I.H., ExtramuralPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20060417PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (Bone Morphogenetic Proteins)RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Bone Morphogenetic Proteins/geneticsMH - Chiroptera/anatomy & histology/embryology/*genetics/*physiologyMH - DNA, Complementary/geneticsMH - *Evolution, MolecularMH - *Flight, AnimalMH - FossilsMH - MiceMH - Molecular Sequence DataMH - Signal TransductionMH - Wing/*anatomy & histology/embryology/*physiologyPMC - PMC1458926OID - NLM: PMC1458926EDAT- 2006/04/19 09:00MHDA- 2006/06/16 09:00CRDT- 2006/04/19 09:00PHST- 2006/04/17 [aheadofprint]AID - 0509716103 [pii]AID - 10.1073/pnas.0509716103 [doi]PST - ppublishSO - Proc Natl Acad Sci U S A. 2006 Apr 25;103(17):6581-6. Epub 2006 Apr 17.

PMID- 11353869OWN - NLMSTAT- MEDLINEDA - 20010524DCOM- 20010719LR - 20081120IS - 0027-8424 (Print)VI - 98IP - 11DP - 2001 May 22TI - Integrated fossil and molecular data reconstruct bat echolocation.PG - 6241-6AB - Molecular and morphological data have important roles in illuminating evolutionary history. DNA data often yield well resolved phylogenies for living taxa, but are generally unattainable for fossils. A distinct advantage of morphology is that some types of morphological data may be collected for extinct and extant taxa. Fossils provide a unique window on evolutionary history and may preserve combinations of primitive and derived characters that are not found in extant taxa. Given their unique character complexes, fossils are critical in documenting sequences of character transformation over geologic time and may

elucidate otherwise ambiguous patterns of evolution that are not revealed by molecular data alone. Here, we employ a methodological approach that allows for the integration of molecular and paleontological data in deciphering one of the most innovative features in the evolutionary history of mammals-laryngeal echolocation in bats. Molecular data alone, including an expanded data set that includes new sequences for the A2AB gene, suggest that microbats are paraphyletic but do not resolve whether laryngeal echolocation evolved independently in different microbat lineages or evolved in the common ancestor of bats and was subsequently lost in megabats. When scaffolds from molecular phylogenies are incorporated into parsimony analyses of morphological characters, including morphological characters for the Eocene taxa Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx, the resulting trees suggest that laryngeal echolocation evolved in the common ancestor of fossil and extant bats and was subsequently lost in megabats. Molecular dating suggests that crown-group bats last shared a common ancestor 52 to 54 million years ago.AD - Department of Biology, University of California, Riverside, CA 92521, USA.FAU - Springer, M SAU - Springer MSFAU - Teeling, E CAU - Teeling ECFAU - Madsen, OAU - Madsen OFAU - Stanhope, M JAU - Stanhope MJFAU - de Jong, W WAU - de Jong WWLA - engSI - GENBANK/AF337537SI - GENBANK/AF337538SI - GENBANK/AF337539SI - GENBANK/AF337540SI - GENBANK/AF337541SI - GENBANK/AF337542SI - GENBANK/AF337543PT - Journal ArticlePT - Research Support, Non-U.S. Gov'tPT - Research SuppoPMID- 18198331OWN - NLMSTAT- MEDLINEDA - 20080116DCOM- 20080313LR - 20081121IS - 0890-9369 (Print)VI - 22IP - 2DP - 2008 Jan 15TI - Understanding of bat wing evolution takes flight.

PG - 121-4AD - Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.FAU - Cooper, Kimberly LAU - Cooper KLFAU - Tabin, Clifford JAU - Tabin CJLA - engGR - F32 HD 052349/HD/NICHD NIH HHS/United StatesGR - R37 HD 32443/HD/NICHD NIH HHS/United StatesPT - CommentPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPL - United StatesTA - Genes DevJT - Genes & developmentJID - 8711660RN - 0 (Homeodomain Proteins)SB - IMCON - Genes Dev. 2008 Jan 15;22(2):141-51. PMID: 18198333MH - AnimalsMH - Chiroptera/*geneticsMH - *EvolutionMH - Forelimb/anatomy & histologyMH - FossilsMH - *Genetic VariationMH - Homeodomain Proteins/*geneticsMH - Wing/*growth & developmentEDAT- 2008/01/17 09:00MHDA- 2008/03/14 09:00CRDT- 2008/01/17 09:00AID - 22/2/121 [pii]AID - 10.1101/gad.1639108 [doi]PST - ppublishSO - Genes Dev. 2008 Jan 15;22(2):121-4.

PMID- 16618938OWN - NLMSTAT- MEDLINEDA - 20060427DCOM- 20060615LR - 20081120IS - 0027-8424 (Print)VI - 103IP - 17DP - 2006 Apr 25TI - Development of bat flight: morphologic and molecular evolution of bat wing digits.PG - 6581-6AB - The earliest fossil bats resemble their modern counterparts in possessing greatly elongated digits to support the wing membrane, which is an anatomical hallmark of powered flight. To quantitatively confirm these similarities, we performed a morphometric analysis of wing bones from fossil and modern bats. We found that the lengths of the third, fourth, and fifth digits (the primary supportive elements of the wing) have remained constant relative to body size over the last 50 million years. This absence of transitional forms in the fossil record

led us to look elsewhere to understand bat wing evolution. Investigating embryonic development, we found that the digits in bats (Carollia perspicillata) are initially similar in size to those of mice (Mus musculus) but that, subsequently, bat digits greatly lengthen. The developmental timing of the change in wing digit length points to a change in longitudinal cartilage growth, a process that depends on the relative proliferation and differentiation of chondrocytes. We found that bat forelimb digits exhibit relatively high rates of chondrocyte proliferation and differentiation. We show that bone morphogenetic protein 2 (Bmp2) can stimulate cartilage proliferation and differentiation and increase digit length in the bat embryonic forelimb. Also, we show that Bmp2 expression and Bmp signaling are increased in bat forelimb embryonic digits relative to mouse or bat hind limb digits. Together, our results suggest that an up-regulation of the Bmp pathway is one of the major factors in the developmental elongation of bat forelimb digits, and it is potentially a key mechanism in their evolutionary elongation as well.AD - Howard Hughes Medical Institute, Department of Pediatrics, Section of Developmental Biology, University of Colorado at Denver and Health Sciences Center, 12800 East 19th Avenue, Aurora, CO 80045, USA.FAU - Sears, Karen EAU - Sears KEFAU - Behringer, Richard RAU - Behringer RRFAU - Rasweiler, John J 4thAU - Rasweiler JJ 4thFAU - Niswander, Lee AAU - Niswander LALA - engSI - GENBANK/DQ279782SI - GENBANK/DQ279783SI - GENBANK/DQ279784SI - GENBANK/DQ279785GR - F32 HD050042-01/HD/NICHD NIH HHS/United StatesGR - HD32427/HD/NICHD NIH HHS/United StatesPT - Comparative StudyPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20060417PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (Bone Morphogenetic Proteins)RN - 0 (DNA, Complementary)SB - IMMH - Animals

MH - Base SequenceMH - Bone Morphogenetic Proteins/geneticsMH - Chiroptera/anatomy & histology/embryology/*genetics/*physiologyMH - DNA, Complementary/geneticsMH - *Evolution, MolecularMH - *Flight, AnimalMH - FossilsMH - MiceMH - Molecular Sequence DataMH - Signal TransductionMH - Wing/*anatomy & histology/embryology/*physiologyPMC - PMC1458926OID - NLM: PMC1458926EDAT- 2006/04/19 09:00MHDA- 2006/06/16 09:00CRDT- 2006/04/19 09:00PHST- 2006/04/17 [aheadofprint]AID - 0509716103 [pii]AID - 10.1073/pnas.0509716103 [doi]PST - ppublishSO - Proc Natl Acad Sci U S A. 2006 Apr 25;103(17):6581-6. Epub 2006 Apr 17.

PMID- 11353869OWN - NLMSTAT- MEDLINEDA - 20010524DCOM- 20010719LR - 20081120IS - 0027-8424 (Print)VI - 98IP - 11DP - 2001 May 22TI - Integrated fossil and molecular data reconstruct bat echolocation.PG - 6241-6AB - Molecular and morphological data have important roles in illuminating evolutionary history. DNA data often yield well resolved phylogenies for living taxa, but are generally unattainable for fossils. A distinct advantage of morphology is that some types of morphological data may be collected for extinct and extant taxa. Fossils provide a unique window on evolutionary history and may preserve combinations of primitive and derived characters that are not found in extant taxa. Given their unique character complexes, fossils are critical in documenting sequences of character transformation over geologic time and may elucidate otherwise ambiguous patterns of evolution that are not revealed by molecular data alone. Here, we employ a methodological approach that allows for the integration of molecular and paleontological data in deciphering one of the most innovative features in the evolutionary history of mammals-laryngeal echolocation in bats. Molecular data alone, including an expanded data set that includes new sequences for the A2AB gene, suggest that microbats are paraphyletic but do not resolve whether laryngeal echolocation evolved independently in

different microbat lineages or evolved in the common ancestor of bats and was subsequently lost in megabats. When scaffolds from molecular phylogenies are incorporated into parsimony analyses of morphological characters, including morphological characters for the Eocene taxa Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx, the resulting trees suggest that laryngeal echolocation evolved in the common ancestor of fossil and extant bats and was subsequently lost in megabats. Molecular dating suggests that crown-group bats last shared a common ancestor 52 to 54 million years ago.AD - Department of Biology, University of California, Riverside, CA 92521, USA.FAU - Springer, M SAU - Springer MSFAU - Teeling, E CAU - Teeling ECFAU - Madsen, OAU - Madsen OFAU - Stanhope, M JAU - Stanhope MJFAU - de Jong, W WAU - de Jong WWLA - engSI - GENBANK/AF337537SI - GENBANK/AF337538SI - GENBANK/AF337539SI - GENBANK/AF337540SI - GENBANK/AF337541SI - GENBANK/AFPMID- 18198331OWN - NLMSTAT- MEDLINEDA - 20080116DCOM- 20080313LR - 20081121IS - 0890-9369 (Print)VI - 22IP - 2DP - 2008 Jan 15TI - Understanding of bat wing evolution takes flight.PG - 121-4AD - Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.FAU - Cooper, Kimberly LAU - Cooper KLFAU - Tabin, Clifford JAU - Tabin CJLA - engGR - F32 HD 052349/HD/NICHD NIH HHS/United StatesGR - R37 HD 32443/HD/NICHD NIH HHS/United StatesPT - CommentPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPL - United StatesTA - Genes DevJT - Genes & developmentJID - 8711660

RN - 0 (Homeodomain Proteins)SB - IMCON - Genes Dev. 2008 Jan 15;22(2):141-51. PMID: 18198333MH - AnimalsMH - Chiroptera/*geneticsMH - *EvolutionMH - Forelimb/anatomy & histologyMH - FossilsMH - *Genetic VariationMH - Homeodomain Proteins/*geneticsMH - Wing/*growth & developmentEDAT- 2008/01/17 09:00MHDA- 2008/03/14 09:00CRDT- 2008/01/17 09:00AID - 22/2/121 [pii]AID - 10.1101/gad.1639108 [doi]PST - ppublishSO - Genes Dev. 2008 Jan 15;22(2):121-4.

PMID- 16618938OWN - NLMSTAT- MEDLINEDA - 20060427DCOM- 20060615LR - 20081120IS - 0027-8424 (Print)VI - 103IP - 17DP - 2006 Apr 25TI - Development of bat flight: morphologic and molecular evolution of bat wing digits.PG - 6581-6AB - The earliest fossil bats resemble their modern counterparts in possessing greatly elongated digits to support the wing membrane, which is an anatomical hallmark of powered flight. To quantitatively confirm these similarities, we performed a morphometric analysis of wing bones from fossil and modern bats. We found that the lengths of the third, fourth, and fifth digits (the primary supportive elements of the wing) have remained constant relative to body size over the last 50 million years. This absence of transitional forms in the fossil record led us to look elsewhere to understand bat wing evolution. Investigating embryonic development, we found that the digits in bats (Carollia perspicillata) are initially similar in size to those of mice (Mus musculus) but that, subsequently, bat digits greatly lengthen. The developmental timing of the change in wing digit length points to a change in longitudinal cartilage growth, a process that depends on the relative proliferation and differentiation of chondrocytes. We found that bat forelimb digits exhibit relatively high rates of chondrocyte proliferation and differentiation. We show that bone morphogenetic protein 2 (Bmp2) can stimulate cartilage proliferation and differentiation and incre

ase digit length in the bat embryonic forelimb. Also, we show that Bmp2 expression and Bmp signaling are increased in bat forelimb embryonic digits relative to mouse or bat hind limb digits. Together, our results suggest that an up-regulation of the Bmp pathway is one of the major factors in the developmental elongation of bat forelimb digits, and it is potentially a key mechanism in their evolutionary elongation as well.AD - Howard Hughes Medical Institute, Department of Pediatrics, Section of Developmental Biology, University of Colorado at Denver and Health Sciences Center, 12800 East 19th Avenue, Aurora, CO 80045, USA.FAU - Sears, Karen EAU - Sears KEFAU - Behringer, Richard RAU - Behringer RRFAU - Rasweiler, John J 4thAU - Rasweiler JJ 4thFAU - Niswander, Lee AAU - Niswander LALA - engSI - GENBANK/DQ279782SI - GENBANK/DQ279783SI - GENBANK/DQ279784SI - GENBANK/DQ279785GR - F32 HD050042-01/HD/NICHD NIH HHS/United StatesGR - HD32427/HD/NICHD NIH HHS/United StatesPT - Comparative StudyPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20060417PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (Bone Morphogenetic Proteins)RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Bone Morphogenetic Proteins/geneticsMH - Chiroptera/anatomy & histology/embryology/*genetics/*physiologyMH - DNA, Complementary/geneticsMH - *Evolution, MolecularMH - *Flight, AnimalMH - FossilsMH - MiceMH - Molecular Sequence DataMH - Signal TransductionMH - Wing/*anatomy & histology/embryology/*physiologyPMC - PMC1458926OID - NLM: PMC1458926EDAT- 2006/04/19 09:00MHDA- 2006/06/16 09:00CRDT- 2006/04/19 09:00

PHST- 2006/04/17 [aheadofprint]AID - 0509716103 [pii]AID - 10.1073/pnas.0509716103 [doi]PST - ppublishSO - Proc Natl Acad Sci U S A. 2006 Apr 25;103(17):6581-6. Epub 2006 Apr 17.

PMID- 11353869OWN - NLMSTAT- MEDLINEDA - 20010524DCOM- 20010719LR - 20081120IS - 0027-8424 (Print)VI - 98IP - 11DP - 2001 May 22TI - Integrated fossil and molecular data reconstruct bat echolocation.PG - 6241-6AB - Molecular and morphological data have important roles in illuminating evolutionary history. DNA data often yield well resolved phylogenies for living taxa, but are generally unattainable for fossils. A distinct advantage of morphology is that some types of morphological data may be collected for extinct and extant taxa. Fossils provide a unique window on evolutionary history and may preserve combinations of primitive and derived characters that are not found in extant taxa. Given their unique character complexes, fossils are critical in documenting sequences of character transformation over geologic time and may elucidate otherwise ambiguous patterns of evolution that are not revealed by molecular data alone. Here, we employ a methodological approach that allows for the integration of molecular and paleontological data in deciphering one of the most innovative features in the evolutionary history of mammals-laryngeal echolocation in bats. Molecular data alone, including an expanded data set that includes new sequences for the A2AB gene, suggest that microbats are paraphyletic but do not resolve whether laryngeal echolocation evolved independently in different microbat lineages or evolved in the common ancestor of bats and was subsequently lost in megabats. When scaffolds from molecular phylogenies are incorporated into parsimony analyses of morphological characters, including morphological characters for the Eocene taxa Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx, the resulting trees suggest that laryngeal echolocation evolved in the common ancestor of fossil and extant bats and was subsequently lost in megabats. Molecular dating suggests that crown-group bats last shared a common ancestor 52 to 54 million years ago.AD - Department of Biology, University of California, Riverside, CA 92521, USA.

FAU - Springer, M SAU - Springer MSFAU - Teeling, E CAU - Teeling ECFAU - Madsen, OAU - Madsen OFAU - Stanhope, M JAU - Stanhope MJFAU - de Jong, W WAU - de Jong WWLA - engSI - GENBANK/AF337537SI - GENBANK/AF337538SI - GENBANK/AF337539SI - GENBANK/PMID- 18198331OWN - NLMSTAT- MEDLINEDA - 20080116DCOM- 20080313LR - 20081121IS - 0890-9369 (Print)VI - 22IP - 2DP - 2008 Jan 15TI - Understanding of bat wing evolution takes flight.PG - 121-4AD - Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.FAU - Cooper, Kimberly LAU - Cooper KLFAU - Tabin, Clifford JAU - Tabin CJLA - engGR - F32 HD 052349/HD/NICHD NIH HHS/United StatesGR - R37 HD 32443/HD/NICHD NIH HHS/United StatesPT - CommentPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPL - United StatesTA - Genes DevJT - Genes & developmentJID - 8711660RN - 0 (Homeodomain Proteins)SB - IMCON - Genes Dev. 2008 Jan 15;22(2):141-51. PMID: 18198333MH - AnimalsMH - Chiroptera/*geneticsMH - *EvolutionMH - Forelimb/anatomy & histologyMH - FossilsMH - *Genetic VariationMH - Homeodomain Proteins/*geneticsMH - Wing/*growth & developmentEDAT- 2008/01/17 09:00MHDA- 2008/03/14 09:00CRDT- 2008/01/17 09:00AID - 22/2/121 [pii]AID - 10.1101/gad.1639108 [doi]PST - ppublishSO - Genes Dev. 2008 Jan 15;22(2):121-4.

PMID- 16618938OWN - NLMSTAT- MEDLINEDA - 20060427DCOM- 20060615LR - 20081120IS - 0027-8424 (Print)VI - 103IP - 17DP - 2006 Apr 25TI - Development of bat flight: morphologic and molecular evolution of bat wing digits.PG - 6581-6AB - The earliest fossil bats resemble their modern counterparts in possessing greatly elongated digits to support the wing membrane, which is an anatomical hallmark of powered flight. To quantitatively confirm these similarities, we performed a morphometric analysis of wing bones from fossil and modern bats. We found that the lengths of the third, fourth, and fifth digits (the primary supportive elements of the wing) have remained constant relative to body size over the last 50 million years. This absence of transitional forms in the fossil record led us to look elsewhere to understand bat wing evolution. Investigating embryonic development, we found that the digits in bats (Carollia perspicillata) are initially similar in size to those of mice (Mus musculus) but that, subsequently, bat digits greatly lengthen. The developmental timing of the change in wing digit length points to a change in longitudinal cartilage growth, a process that depends on the relative proliferation and differentiation of chondrocytes. We found that bat forelimb digits exhibit relatively high rates of chondrocyte proliferation and differentiation. We show that bone morphogenetic protein 2 (Bmp2) can stimulate cartilage proliferation and differentiation and increase digit length in the bat embryonic forelimb. Also, we show that Bmp2 expression and Bmp signaling are increased in bat forelimb embryonic digits relative to mouse or bat hind limb digits. Together, our results suggest that an up-regulation of the Bmp pathway is one of the major factors in the developmental elongation of bat forelimb digits, and it is potentially a key mechanism in their evolutionary elongation as well.AD - Howard Hughes Medical Institute, Department of Pediatrics, Section of Developmental Biology, University of Colorado at Denver and Health Sciences Center, 12800 East 19th Avenue, Aurora, CO 80045, USA.FAU - Sears, Karen EAU - Sears KEFAU - Behringer, Richard R

AU - Behringer RRFAU - Rasweiler, John J 4thAU - Rasweiler JJ 4thFAU - Niswander, Lee AAU - Niswander LALA - engSI - GENBANK/DQ279782SI - GENBANK/DQ279783SI - GENBANK/DQ279784SI - GENBANK/DQ279785GR - F32 HD050042-01/HD/NICHD NIH HHS/United StatesGR - HD32427/HD/NICHD NIH HHS/United StatesPT - Comparative StudyPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20060417PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (Bone Morphogenetic Proteins)RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Bone Morphogenetic Proteins/geneticsMH - Chiroptera/anatomy & histology/embryology/*genetics/*physiologyMH - DNA, Complementary/geneticsMH - *Evolution, MolecularMH - *Flight, AnimalMH - FossilsMH - MiceMH - Molecular Sequence DataMH - Signal TransductionMH - Wing/*anatomy & histology/embryology/*physiologyPMC - PMC1458926OID - NLM: PMC1458926EDAT- 2006/04/19 09:00MHDA- 2006/06/16 09:00CRDT- 2006/04/19 09:00PHST- 2006/04/17 [aheadofprint]AID - 0509716103 [pii]AID - 10.1073/pnas.0509716103 [doi]PST - ppublishSO - Proc Natl Acad Sci U S A. 2006 Apr 25;103(17):6581-6. Epub 2006 Apr 17.

PMID- 11353869OWN - NLMSTAT- MEDLINEDA - 20010524DCOM- 20010719LR - 20081120IS - 0027-8424 (Print)VI - 98IP - 11DP - 2001 May 22TI - Integrated fossil and molecular data reconstruct bat echolocation.PG - 6241-6

AB - Molecular and morphological data have important roles in illuminating evolutionary history. DNA data often yield well resolved phylogenies for living taxa, but are generally unattainable for fossils. A distinct advantage of morphology is that some types of morphological data may be collected for extinct and extant taxa. Fossils provide a unique window on evolutionary history and may preserve combinations of primitive and derived characters that are not found in extant taxa. Given their unique character complexes, fossils are critical in documenting sequences of character transformation over geologic time and may elucidate otherwise ambiguous patterns of evolution that are not revealed by molecular data alone. Here, we employ a methodological approach that allows for the integration of molecular and paleontological data in deciphering one of the most innovative features in the evolutionary history of mammals-laryngeal echolocation in bats. Molecular data alone, including an expanded data set that includes new sequences for the A2AB gene, suggest that microbats are paraphyletic but do not resolve whether laryngeal echolocation evolved independently in different microbat lineages or evolved in the common ancestor of bats and was subsequently lost in megabats. When scaffolds from molecular phylogenies are incorporated into parsimony analyses of morphological characters, including morphological characters for the Eocene taxa Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx, the resulting trees suggest that laryngeal echolocation evolved in the common ancestor of fossil and extant bats and was subsequently lost in megabats. Molecular dating suggests that crown-group bats last shared a common ancestor 52 to 54 million years ago.AD - Department of Biology, University of California, Riverside, CA 92521, USA.FAU - Springer, M SAU - Springer MSFAU - Teeling, E CAU - Teeling ECFAU - Madsen, OAU - Madsen OFAU - Stanhope, M JAU - Stanhope MJFAU - de Jong, W WAU - de Jong WPMID- 18198331OWN - NLMSTAT- MEDLINEDA - 20080116DCOM- 20080313LR - 20081121IS - 0890-9369 (Print)VI - 22

IP - 2DP - 2008 Jan 15TI - Understanding of bat wing evolution takes flight.PG - 121-4AD - Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.FAU - Cooper, Kimberly LAU - Cooper KLFAU - Tabin, Clifford JAU - Tabin CJLA - engGR - F32 HD 052349/HD/NICHD NIH HHS/United StatesGR - R37 HD 32443/HD/NICHD NIH HHS/United StatesPT - CommentPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPL - United StatesTA - Genes DevJT - Genes & developmentJID - 8711660RN - 0 (Homeodomain Proteins)SB - IMCON - Genes Dev. 2008 Jan 15;22(2):141-51. PMID: 18198333MH - AnimalsMH - Chiroptera/*geneticsMH - *EvolutionMH - Forelimb/anatomy & histologyMH - FossilsMH - *Genetic VariationMH - Homeodomain Proteins/*geneticsMH - Wing/*growth & developmentEDAT- 2008/01/17 09:00MHDA- 2008/03/14 09:00CRDT- 2008/01/17 09:00AID - 22/2/121 [pii]AID - 10.1101/gad.1639108 [doi]PST - ppublishSO - Genes Dev. 2008 Jan 15;22(2):121-4.

PMID- 16618938OWN - NLMSTAT- MEDLINEDA - 20060427DCOM- 20060615LR - 20081120IS - 0027-8424 (Print)VI - 103IP - 17DP - 2006 Apr 25TI - Development of bat flight: morphologic and molecular evolution of bat wing digits.PG - 6581-6AB - The earliest fossil bats resemble their modern counterparts in possessing greatly elongated digits to support the wing membrane, which is an anatomical hallmark of powered flight. To quantitatively confirm these similarities, we performed a morphometric analysis of wing bones from fossil and modern bats. We found that the lengths of the third, fourth, and fifth digits (the primary supportive

elements of the wing) have remained constant relative to body size over the last 50 million years. This absence of transitional forms in the fossil record led us to look elsewhere to understand bat wing evolution. Investigating embryonic development, we found that the digits in bats (Carollia perspicillata) are initially similar in size to those of mice (Mus musculus) but that, subsequently, bat digits greatly lengthen. The developmental timing of the change in wing digit length points to a change in longitudinal cartilage growth, a process that depends on the relative proliferation and differentiation of chondrocytes. We found that bat forelimb digits exhibit relatively high rates of chondrocyte proliferation and differentiation. We show that bone morphogenetic protein 2 (Bmp2) can stimulate cartilage proliferation and differentiation and increase digit length in the bat embryonic forelimb. Also, we show that Bmp2 expression and Bmp signaling are increased in bat forelimb embryonic digits relative to mouse or bat hind limb digits. Together, our results suggest that an up-regulation of the Bmp pathway is one of the major factors in the developmental elongation of bat forelimb digits, and it is potentially a key mechanism in their evolutionary elongation as well.AD - Howard Hughes Medical Institute, Department of Pediatrics, Section of Developmental Biology, University of Colorado at Denver and Health Sciences Center, 12800 East 19th Avenue, Aurora, CO 80045, USA.FAU - Sears, Karen EAU - Sears KEFAU - Behringer, Richard RAU - Behringer RRFAU - Rasweiler, John J 4thAU - Rasweiler JJ 4thFAU - Niswander, Lee AAU - Niswander LALA - engSI - GENBANK/DQ279782SI - GENBANK/DQ279783SI - GENBANK/DQ279784SI - GENBANK/DQ279785GR - F32 HD050042-01/HD/NICHD NIH HHS/United StatesGR - HD32427/HD/NICHD NIH HHS/United StatesPT - Comparative StudyPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20060417PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (Bone Morphogenetic Proteins)

RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Bone Morphogenetic Proteins/geneticsMH - Chiroptera/anatomy & histology/embryology/*genetics/*physiologyMH - DNA, Complementary/geneticsMH - *Evolution, MolecularMH - *Flight, AnimalMH - FossilsMH - MiceMH - Molecular Sequence DataMH - Signal TransductionMH - Wing/*anatomy & histology/embryology/*physiologyPMC - PMC1458926OID - NLM: PMC1458926EDAT- 2006/04/19 09:00MHDA- 2006/06/16 09:00CRDT- 2006/04/19 09:00PHST- 2006/04/17 [aheadofprint]AID - 0509716103 [pii]AID - 10.1073/pnas.0509716103 [doi]PST - ppublishSO - Proc Natl Acad Sci U S A. 2006 Apr 25;103(17):6581-6. Epub 2006 Apr 17.

PMID- 11353869OWN - NLMSTAT- MEDLINEDA - 20010524DCOM- 20010719LR - 20081120IS - 0027-8424 (Print)VI - 98IP - 11DP - 2001 May 22TI - Integrated fossil and molecular data reconstruct bat echolocation.PG - 6241-6AB - Molecular and morphological data have important roles in illuminating evolutionary history. DNA data often yield well resolved phylogenies for living taxa, but are generally unattainable for fossils. A distinct advantage of morphology is that some types of morphological data may be collected for extinct and extant taxa. Fossils provide a unique window on evolutionary history and may preserve combinations of primitive and derived characters that are not found in extant taxa. Given their unique character complexes, fossils are critical in documenting sequences of character transformation over geologic time and may elucidate otherwise ambiguous patterns of evolution that are not revealed by molecular data alone. Here, we employ a methodological approach that allows for the integration of molecular and paleontological data in deciphering one of the most innovative features in the evolutionary history of mammals-laryngeal echolocation in bats. Molecular data alone, including an expanded data set that

includes new sequences for the A2AB gene, suggest that microbats are paraphyletic but do not resolve whether laryngeal echolocation evolved independently in different microbat lineages or evolved in the common ancestor of bats and was subsequently lost in megabats. When scaffolds from molecular phylogenies are incorporated into parsimony analyses of morphological characters, including morphological characters for the Eocene taxa Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx, the resulting trees suggest that laryngeal echolocation evolved in the common ancestor of fossil and extant bats and was subsequently lost in megabats. Molecular dating suggests that crown-group bats last shared a common ancestor 52 to 54 million years ago.AD - Department of Biology, University of California, Riverside, CA 92521, USA.FAU - Springer, M SAU - Springer MSFAU - Teeling, E CAU - Teeling ECFAU - Madsen, OAU - Madsen OFAU - Stanhope, M JAU - Stanhope MJFAU - de Jong, W WAU - de Jong WWLA - engSI - GENBANK/AF337537SI - GENBANK/AF337538SI - GENBANK/AF337539SI - GENBANK/AF337540SI - GENBANK/AF337541SI - GENBANK/AF337542SI - GENBANK/AF337543PT - Journal ArticlePT - Research Support, Non-U.S. Gov'tPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20010515PL - United StatesPMID- 18198331OWN - NLMSTAT- MEDLINEDA - 20080116DCOM- 20080313LR - 20081121IS - 0890-9369 (Print)VI - 22IP - 2DP - 2008 Jan 15TI - Understanding of bat wing evolution takes flight.PG - 121-4AD - Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.FAU - Cooper, Kimberly LAU - Cooper KLFAU - Tabin, Clifford JAU - Tabin CJLA - eng

GR - F32 HD 052349/HD/NICHD NIH HHS/United StatesGR - R37 HD 32443/HD/NICHD NIH HHS/United StatesPT - CommentPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPL - United StatesTA - Genes DevJT - Genes & developmentJID - 8711660RN - 0 (Homeodomain Proteins)SB - IMCON - Genes Dev. 2008 Jan 15;22(2):141-51. PMID: 18198333MH - AnimalsMH - Chiroptera/*geneticsMH - *EvolutionMH - Forelimb/anatomy & histologyMH - FossilsMH - *Genetic VariationMH - Homeodomain Proteins/*geneticsMH - Wing/*growth & developmentEDAT- 2008/01/17 09:00MHDA- 2008/03/14 09:00CRDT- 2008/01/17 09:00AID - 22/2/121 [pii]AID - 10.1101/gad.1639108 [doi]PST - ppublishSO - Genes Dev. 2008 Jan 15;22(2):121-4.

PMID- 16618938OWN - NLMSTAT- MEDLINEDA - 20060427DCOM- 20060615LR - 20081120IS - 0027-8424 (Print)VI - 103IP - 17DP - 2006 Apr 25TI - Development of bat flight: morphologic and molecular evolution of bat wing digits.PG - 6581-6AB - The earliest fossil bats resemble their modern counterparts in possessing greatly elongated digits to support the wing membrane, which is an anatomical hallmark of powered flight. To quantitatively confirm these similarities, we performed a morphometric analysis of wing bones from fossil and modern bats. We found that the lengths of the third, fourth, and fifth digits (the primary supportive elements of the wing) have remained constant relative to body size over the last 50 million years. This absence of transitional forms in the fossil record led us to look elsewhere to understand bat wing evolution. Investigating embryonic development, we found that the digits in bats (Carollia perspicillata) are initially similar in size to those of mice (Mus musculus) but that, subsequently, bat digits greatly lengthen. The developmental timing of the change in win

g digit length points to a change in longitudinal cartilage growth, a process that depends on the relative proliferation and differentiation of chondrocytes. We found that bat forelimb digits exhibit relatively high rates of chondrocyte proliferation and differentiation. We show that bone morphogenetic protein 2 (Bmp2) can stimulate cartilage proliferation and differentiation and increase digit length in the bat embryonic forelimb. Also, we show that Bmp2 expression and Bmp signaling are increased in bat forelimb embryonic digits relative to mouse or bat hind limb digits. Together, our results suggest that an up-regulation of the Bmp pathway is one of the major factors in the developmental elongation of bat forelimb digits, and it is potentially a key mechanism in their evolutionary elongation as well.AD - Howard Hughes Medical Institute, Department of Pediatrics, Section of Developmental Biology, University of Colorado at Denver and Health Sciences Center, 12800 East 19th Avenue, Aurora, CO 80045, USA.FAU - Sears, Karen EAU - Sears KEFAU - Behringer, Richard RAU - Behringer RRFAU - Rasweiler, John J 4thAU - Rasweiler JJ 4thFAU - Niswander, Lee AAU - Niswander LALA - engSI - GENBANK/DQ279782SI - GENBANK/DQ279783SI - GENBANK/DQ279784SI - GENBANK/DQ279785GR - F32 HD050042-01/HD/NICHD NIH HHS/United StatesGR - HD32427/HD/NICHD NIH HHS/United StatesPT - Comparative StudyPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20060417PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (Bone Morphogenetic Proteins)RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Bone Morphogenetic Proteins/geneticsMH - Chiroptera/anatomy & histology/embryology/*genetics/*physiologyMH - DNA, Complementary/geneticsMH - *Evolution, MolecularMH - *Flight, AnimalMH - Fossils

MH - MiceMH - Molecular Sequence DataMH - Signal TransductionMH - Wing/*anatomy & histology/embryology/*physiologyPMC - PMC1458926OID - NLM: PMC1458926EDAT- 2006/04/19 09:00MHDA- 2006/06/16 09:00CRDT- 2006/04/19 09:00PHST- 2006/04/17 [aheadofprint]AID - 0509716103 [pii]AID - 10.1073/pnas.0509716103 [doi]PST - ppublishSO - Proc Natl Acad Sci U S A. 2006 Apr 25;103(17):6581-6. Epub 2006 Apr 17.

PMID- 11353869OWN - NLMSTAT- MEDLINEDA - 20010524DCOM- 20010719LR - 20081120IS - 0027-8424 (Print)VI - 98IP - 11DP - 2001 May 22TI - Integrated fossil and molecular data reconstruct bat echolocation.PG - 6241-6AB - Molecular and morphological data have important roles in illuminating evolutionary history. DNA data often yield well resolved phylogenies for living taxa, but are generally unattainable for fossils. A distinct advantage of morphology is that some types of morphological data may be collected for extinct and extant taxa. Fossils provide a unique window on evolutionary history and may preserve combinations of primitive and derived characters that are not found in extant taxa. Given their unique character complexes, fossils are critical in documenting sequences of character transformation over geologic time and may elucidate otherwise ambiguous patterns of evolution that are not revealed by molecular data alone. Here, we employ a methodological approach that allows for the integration of molecular and paleontological data in deciphering one of the most innovative features in the evolutionary history of mammals-laryngeal echolocation in bats. Molecular data alone, including an expanded data set that includes new sequences for the A2AB gene, suggest that microbats are paraphyletic but do not resolve whether laryngeal echolocation evolved independently in different microbat lineages or evolved in the common ancestor of bats and was subsequently lost in megabats. When scaffolds from molecular phylogenies are incorporated into parsimony analyses of morphological characters, including morphological characters for the Eocene taxa Icaronycteris, Archaeonycteri

s, Hassianycteris, and Palaeochiropteryx, the resulting trees suggest that laryngeal echolocation evolved in the common ancestor of fossil and extant bats and was subsequently lost in megabats. Molecular dating suggests that crown-group bats last shared a common ancestor 52 to 54 million years ago.AD - Department of Biology, University of California, Riverside, CA 92521, USA.FAU - Springer, M SAU - Springer MSFAU - Teeling, E CAU - Teeling ECFAU - Madsen, OAU - Madsen OFAU - Stanhope, M JAU - Stanhope MJFAU - de Jong, W WAU - de Jong WWLA - engSI - GENBANK/AF337537SI - GENBANK/AF337538SI - GENBANK/AF337539SI - GENBANK/AF337540SI - GENBANK/AF337541SI - GENBANK/AF337542SI - GENBANK/AF337543PT - Journal ArticlePT - Research Support, Non-U.S. Gov'tPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20010515PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequencePMID- 18198331OWN - NLMSTAT- MEDLINEDA - 20080116DCOM- 20080313LR - 20081121IS - 0890-9369 (Print)VI - 22IP - 2DP - 2008 Jan 15TI - Understanding of bat wing evolution takes flight.PG - 121-4AD - Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.FAU - Cooper, Kimberly LAU - Cooper KLFAU - Tabin, Clifford JAU - Tabin CJLA - engGR - F32 HD 052349/HD/NICHD NIH HHS/United StatesGR - R37 HD 32443/HD/NICHD NIH HHS/United States

PT - CommentPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPL - United StatesTA - Genes DevJT - Genes & developmentJID - 8711660RN - 0 (Homeodomain Proteins)SB - IMCON - Genes Dev. 2008 Jan 15;22(2):141-51. PMID: 18198333MH - AnimalsMH - Chiroptera/*geneticsMH - *EvolutionMH - Forelimb/anatomy & histologyMH - FossilsMH - *Genetic VariationMH - Homeodomain Proteins/*geneticsMH - Wing/*growth & developmentEDAT- 2008/01/17 09:00MHDA- 2008/03/14 09:00CRDT- 2008/01/17 09:00AID - 22/2/121 [pii]AID - 10.1101/gad.1639108 [doi]PST - ppublishSO - Genes Dev. 2008 Jan 15;22(2):121-4.

PMID- 16618938OWN - NLMSTAT- MEDLINEDA - 20060427DCOM- 20060615LR - 20081120IS - 0027-8424 (Print)VI - 103IP - 17DP - 2006 Apr 25TI - Development of bat flight: morphologic and molecular evolution of bat wing digits.PG - 6581-6AB - The earliest fossil bats resemble their modern counterparts in possessing greatly elongated digits to support the wing membrane, which is an anatomical hallmark of powered flight. To quantitatively confirm these similarities, we performed a morphometric analysis of wing bones from fossil and modern bats. We found that the lengths of the third, fourth, and fifth digits (the primary supportive elements of the wing) have remained constant relative to body size over the last 50 million years. This absence of transitional forms in the fossil record led us to look elsewhere to understand bat wing evolution. Investigating embryonic development, we found that the digits in bats (Carollia perspicillata) are initially similar in size to those of mice (Mus musculus) but that, subsequently, bat digits greatly lengthen. The developmental timing of the change in wing digit length points to a change in longitudinal cartilage growth, a process that

depends on the relative proliferation and differentiation of chondrocytes. We found that bat forelimb digits exhibit relatively high rates of chondrocyte proliferation and differentiation. We show that bone morphogenetic protein 2 (Bmp2) can stimulate cartilage proliferation and differentiation and increase digit length in the bat embryonic forelimb. Also, we show that Bmp2 expression and Bmp signaling are increased in bat forelimb embryonic digits relative to mouse or bat hind limb digits. Together, our results suggest that an up-regulation of the Bmp pathway is one of the major factors in the developmental elongation of bat forelimb digits, and it is potentially a key mechanism in their evolutionary elongation as well.AD - Howard Hughes Medical Institute, Department of Pediatrics, Section of Developmental Biology, University of Colorado at Denver and Health Sciences Center, 12800 East 19th Avenue, Aurora, CO 80045, USA.FAU - Sears, Karen EAU - Sears KEFAU - Behringer, Richard RAU - Behringer RRFAU - Rasweiler, John J 4thAU - Rasweiler JJ 4thFAU - Niswander, Lee AAU - Niswander LALA - engSI - GENBANK/DQ279782SI - GENBANK/DQ279783SI - GENBANK/DQ279784SI - GENBANK/DQ279785GR - F32 HD050042-01/HD/NICHD NIH HHS/United StatesGR - HD32427/HD/NICHD NIH HHS/United StatesPT - Comparative StudyPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20060417PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (Bone Morphogenetic Proteins)RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Bone Morphogenetic Proteins/geneticsMH - Chiroptera/anatomy & histology/embryology/*genetics/*physiologyMH - DNA, Complementary/geneticsMH - *Evolution, MolecularMH - *Flight, AnimalMH - FossilsMH - MiceMH - Molecular Sequence Data

MH - Signal TransductionMH - Wing/*anatomy & histology/embryology/*physiologyPMC - PMC1458926OID - NLM: PMC1458926EDAT- 2006/04/19 09:00MHDA- 2006/06/16 09:00CRDT- 2006/04/19 09:00PHST- 2006/04/17 [aheadofprint]AID - 0509716103 [pii]AID - 10.1073/pnas.0509716103 [doi]PST - ppublishSO - Proc Natl Acad Sci U S A. 2006 Apr 25;103(17):6581-6. Epub 2006 Apr 17.

PMID- 11353869OWN - NLMSTAT- MEDLINEDA - 20010524DCOM- 20010719LR - 20081120IS - 0027-8424 (Print)VI - 98IP - 11DP - 2001 May 22TI - Integrated fossil and molecular data reconstruct bat echolocation.PG - 6241-6AB - Molecular and morphological data have important roles in illuminating evolutionary history. DNA data often yield well resolved phylogenies for living taxa, but are generally unattainable for fossils. A distinct advantage of morphology is that some types of morphological data may be collected for extinct and extant taxa. Fossils provide a unique window on evolutionary history and may preserve combinations of primitive and derived characters that are not found in extant taxa. Given their unique character complexes, fossils are critical in documenting sequences of character transformation over geologic time and may elucidate otherwise ambiguous patterns of evolution that are not revealed by molecular data alone. Here, we employ a methodological approach that allows for the integration of molecular and paleontological data in deciphering one of the most innovative features in the evolutionary history of mammals-laryngeal echolocation in bats. Molecular data alone, including an expanded data set that includes new sequences for the A2AB gene, suggest that microbats are paraphyletic but do not resolve whether laryngeal echolocation evolved independently in different microbat lineages or evolved in the common ancestor of bats and was subsequently lost in megabats. When scaffolds from molecular phylogenies are incorporated into parsimony analyses of morphological characters, including morphological characters for the Eocene taxa Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx, the resulting trees suggest that la

ryngeal echolocation evolved in the common ancestor of fossil and extant bats and was subsequently lost in megabats. Molecular dating suggests that crown-group bats last shared a common ancestor 52 to 54 million years ago.AD - Department of Biology, University of California, Riverside, CA 92521, USA.FAU - Springer, M SAU - Springer MSFAU - Teeling, E CAU - Teeling ECFAU - Madsen, OAU - Madsen OFAU - Stanhope, M JAU - Stanhope MJFAU - de Jong, W WAU - de Jong WWLA - engSI - GENBANK/AF337537SI - GENBANK/AF337538SI - GENBANK/AF337539SI - GENBANK/AF337540SI - GENBANK/AF33PMID- 18198331OWN - NLMSTAT- MEDLINEDA - 20080116DCOM- 20080313LR - 20081121IS - 0890-9369 (Print)VI - 22IP - 2DP - 2008 Jan 15TI - Understanding of bat wing evolution takes flight.PG - 121-4AD - Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.FAU - Cooper, Kimberly LAU - Cooper KLFAU - Tabin, Clifford JAU - Tabin CJLA - engGR - F32 HD 052349/HD/NICHD NIH HHS/United StatesGR - R37 HD 32443/HD/NICHD NIH HHS/United StatesPT - CommentPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPL - United StatesTA - Genes DevJT - Genes & developmentJID - 8711660RN - 0 (Homeodomain Proteins)SB - IMCON - Genes Dev. 2008 Jan 15;22(2):141-51. PMID: 18198333MH - AnimalsMH - Chiroptera/*geneticsMH - *EvolutionMH - Forelimb/anatomy & histologyMH - FossilsMH - *Genetic VariationMH - Homeodomain Proteins/*genetics

MH - Wing/*growth & developmentEDAT- 2008/01/17 09:00MHDA- 2008/03/14 09:00CRDT- 2008/01/17 09:00AID - 22/2/121 [pii]AID - 10.1101/gad.1639108 [doi]PST - ppublishSO - Genes Dev. 2008 Jan 15;22(2):121-4.

PMID- 16618938OWN - NLMSTAT- MEDLINEDA - 20060427DCOM- 20060615LR - 20081120IS - 0027-8424 (Print)VI - 103IP - 17DP - 2006 Apr 25TI - Development of bat flight: morphologic and molecular evolution of bat wing digits.PG - 6581-6AB - The earliest fossil bats resemble their modern counterparts in possessing greatly elongated digits to support the wing membrane, which is an anatomical hallmark of powered flight. To quantitatively confirm these similarities, we performed a morphometric analysis of wing bones from fossil and modern bats. We found that the lengths of the third, fourth, and fifth digits (the primary supportive elements of the wing) have remained constant relative to body size over the last 50 million years. This absence of transitional forms in the fossil record led us to look elsewhere to understand bat wing evolution. Investigating embryonic development, we found that the digits in bats (Carollia perspicillata) are initially similar in size to those of mice (Mus musculus) but that, subsequently, bat digits greatly lengthen. The developmental timing of the change in wing digit length points to a change in longitudinal cartilage growth, a process that depends on the relative proliferation and differentiation of chondrocytes. We found that bat forelimb digits exhibit relatively high rates of chondrocyte proliferation and differentiation. We show that bone morphogenetic protein 2 (Bmp2) can stimulate cartilage proliferation and differentiation and increase digit length in the bat embryonic forelimb. Also, we show that Bmp2 expression and Bmp signaling are increased in bat forelimb embryonic digits relative to mouse or bat hind limb digits. Together, our results suggest that an up-regulation of the Bmp pathway is one of the major factors in the developmental elongation of bat forelimb digits, and it is potentially a key mechanism in their

evolutionary elongation as well.AD - Howard Hughes Medical Institute, Department of Pediatrics, Section of Developmental Biology, University of Colorado at Denver and Health Sciences Center, 12800 East 19th Avenue, Aurora, CO 80045, USA.FAU - Sears, Karen EAU - Sears KEFAU - Behringer, Richard RAU - Behringer RRFAU - Rasweiler, John J 4thAU - Rasweiler JJ 4thFAU - Niswander, Lee AAU - Niswander LALA - engSI - GENBANK/DQ279782SI - GENBANK/DQ279783SI - GENBANK/DQ279784SI - GENBANK/DQ279785GR - F32 HD050042-01/HD/NICHD NIH HHS/United StatesGR - HD32427/HD/NICHD NIH HHS/United StatesPT - Comparative StudyPT - Journal ArticlePT - Research Support, N.I.H., ExtramuralPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20060417PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (Bone Morphogenetic Proteins)RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Bone Morphogenetic Proteins/geneticsMH - Chiroptera/anatomy & histology/embryology/*genetics/*physiologyMH - DNA, Complementary/geneticsMH - *Evolution, MolecularMH - *Flight, AnimalMH - FossilsMH - MiceMH - Molecular Sequence DataMH - Signal TransductionMH - Wing/*anatomy & histology/embryology/*physiologyPMC - PMC1458926OID - NLM: PMC1458926EDAT- 2006/04/19 09:00MHDA- 2006/06/16 09:00CRDT- 2006/04/19 09:00PHST- 2006/04/17 [aheadofprint]AID - 0509716103 [pii]AID - 10.1073/pnas.0509716103 [doi]PST - ppublishSO - Proc Natl Acad Sci U S A. 2006 Apr 25;103(17):6581-6. Epub 2006 Apr 17.

PMID- 11353869OWN - NLMSTAT- MEDLINEDA - 20010524

DCOM- 20010719LR - 20081120IS - 0027-8424 (Print)VI - 98IP - 11DP - 2001 May 22TI - Integrated fossil and molecular data reconstruct bat echolocation.PG - 6241-6AB - Molecular and morphological data have important roles in illuminating evolutionary history. DNA data often yield well resolved phylogenies for living taxa, but are generally unattainable for fossils. A distinct advantage of morphology is that some types of morphological data may be collected for extinct and extant taxa. Fossils provide a unique window on evolutionary history and may preserve combinations of primitive and derived characters that are not found in extant taxa. Given their unique character complexes, fossils are critical in documenting sequences of character transformation over geologic time and may elucidate otherwise ambiguous patterns of evolution that are not revealed by molecular data alone. Here, we employ a methodological approach that allows for the integration of molecular and paleontological data in deciphering one of the most innovative features in the evolutionary history of mammals-laryngeal echolocation in bats. Molecular data alone, including an expanded data set that includes new sequences for the A2AB gene, suggest that microbats are paraphyletic but do not resolve whether laryngeal echolocation evolved independently in different microbat lineages or evolved in the common ancestor of bats and was subsequently lost in megabats. When scaffolds from molecular phylogenies are incorporated into parsimony analyses of morphological characters, including morphological characters for the Eocene taxa Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx, the resulting trees suggest that laryngeal echolocation evolved in the common ancestor of fossil and extant bats and was subsequently lost in megabats. Molecular dating suggests that crown-group bats last shared a common ancestor 52 to 54 million years ago.AD - Department of Biology, University of California, Riverside, CA 92521, USA.FAU - Springer, M SAU - Springer MSFAU - Teeling, E CAU - Teeling ECFAU - Madsen, OAU - Madsen OFAU - Stanhope, M JAU - Stanhope MJFAU - de Jong, W WAU - de Jong WW

LA - engSI - GENBANK/AF337537SI - GENBANK/AF337538SI - GENBANK/AF337539SI - GENBANK/AF337540SI - GENBANK/AF337541SI - GENBANK/AF337542SI - GENBANK/AF337543PT - Journal ArticlePT - Research Support, Non-U.S. Gov'tPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20010515PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Chiroptera/classification/*geneticsMH - DNA, ComplementaryMH - EcosystemMH - *Evolution, MolecularMH - *FossilsMH - HumansMH - Molecular Sequence DataMH - PhylogenyPMC - PMC33452OID - NLM: PMC33452EDAT- 2001/05/17 10:00MHDA- 2001/07/20 10:01CRDT- 2001/05/17 10:00PHST- 2001/05/15 [aheadofprint]AID - 10.1073/pnas.111551998 [doi]AID - 111551998 [pii]PST - ppublishSO - Proc Natl Acad Sci U S A. 2001 May 22;98(11):6241-6. Epub 2001 May 15.7541SI - GENBANK/AF337542SI - GENBANK/AF337543PT - Journal ArticlePT - Research Support, Non-U.S. Gov'tPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20010515PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Chiroptera/classification/*geneticsMH - DNA, ComplementaryMH - EcosystemMH - *Evolution, MolecularMH - *Fossils

MH - HumansMH - Molecular Sequence DataMH - PhylogenyPMC - PMC33452OID - NLM: PMC33452EDAT- 2001/05/17 10:00MHDA- 2001/07/20 10:01CRDT- 2001/05/17 10:00PHST- 2001/05/15 [aheadofprint]AID - 10.1073/pnas.111551998 [doi]AID - 111551998 [pii]PST - ppublishSO - Proc Natl Acad Sci U S A. 2001 May 22;98(11):6241-6. Epub 2001 May 15.

MH - Chiroptera/classification/*geneticsMH - DNA, ComplementaryMH - EcosystemMH - *Evolution, MolecularMH - *FossilsMH - HumansMH - Molecular Sequence DataMH - PhylogenyPMC - PMC33452OID - NLM: PMC33452EDAT- 2001/05/17 10:00MHDA- 2001/07/20 10:01CRDT- 2001/05/17 10:00PHST- 2001/05/15 [aheadofprint]AID - 10.1073/pnas.111551998 [doi]AID - 111551998 [pii]PST - ppublishSO - Proc Natl Acad Sci U S A. 2001 May 22;98(11):6241-6. Epub 2001 May 15.

TA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Chiroptera/classification/*geneticsMH - DNA, ComplementaryMH - EcosystemMH - *Evolution, MolecularMH - *FossilsMH - HumansMH - Molecular Sequence DataMH - PhylogenyPMC - PMC33452OID - NLM: PMC33452EDAT- 2001/05/17 10:00MHDA- 2001/07/20 10:01CRDT- 2001/05/17 10:00PHST- 2001/05/15 [aheadofprint]AID - 10.1073/pnas.111551998 [doi]AID - 111551998 [pii]PST - ppublishSO - Proc Natl Acad Sci U S A. 2001 May 22;98(11):6241-6. Epub 2001 May 15.W

LA - engSI - GENBANK/AF337537SI - GENBANK/AF337538SI - GENBANK/AF337539SI - GENBANK/AF337540SI - GENBANK/AF337541SI - GENBANK/AF337542SI - GENBANK/AF337543PT - Journal ArticlePT - Research Support, Non-U.S. Gov'tPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20010515PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Chiroptera/classification/*geneticsMH - DNA, ComplementaryMH - EcosystemMH - *Evolution, MolecularMH - *FossilsMH - HumansMH - Molecular Sequence DataMH - PhylogenyPMC - PMC33452OID - NLM: PMC33452EDAT- 2001/05/17 10:00MHDA- 2001/07/20 10:01CRDT- 2001/05/17 10:00PHST- 2001/05/15 [aheadofprint]AID - 10.1073/pnas.111551998 [doi]AID - 111551998 [pii]PST - ppublishSO - Proc Natl Acad Sci U S A. 2001 May 22;98(11):6241-6. Epub 2001 May 15.AF337540SI - GENBANK/AF337541SI - GENBANK/AF337542SI - GENBANK/AF337543PT - Journal ArticlePT - Research Support, Non-U.S. Gov'tPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20010515PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Chiroptera/classification/*geneticsMH - DNA, ComplementaryMH - EcosystemMH - *Evolution, Molecular

MH - *FossilsMH - HumansMH - Molecular Sequence DataMH - PhylogenyPMC - PMC33452OID - NLM: PMC33452EDAT- 2001/05/17 10:00MHDA- 2001/07/20 10:01CRDT- 2001/05/17 10:00PHST- 2001/05/15 [aheadofprint]AID - 10.1073/pnas.111551998 [doi]AID - 111551998 [pii]PST - ppublishSO - Proc Natl Acad Sci U S A. 2001 May 22;98(11):6241-6. Epub 2001 May 15.337542SI - GENBANK/AF337543PT - Journal ArticlePT - Research Support, Non-U.S. Gov'tPT - Research Support, U.S. Gov't, Non-P.H.S.DEP - 20010515PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Chiroptera/classification/*geneticsMH - DNA, ComplementaryMH - EcosystemMH - *Evolution, MolecularMH - *FossilsMH - HumansMH - Molecular Sequence DataMH - PhylogenyPMC - PMC33452OID - NLM: PMC33452EDAT- 2001/05/17 10:00MHDA- 2001/07/20 10:01CRDT- 2001/05/17 10:00PHST- 2001/05/15 [aheadofprint]AID - 10.1073/pnas.111551998 [doi]AID - 111551998 [pii]PST - ppublishSO - Proc Natl Acad Sci U S A. 2001 May 22;98(11):6241-6. Epub 2001 May 15.rt, U.S. Gov't, Non-P.H.S.DEP - 20010515PL - United StatesTA - Proc Natl Acad Sci U S AJT - Proceedings of the National Academy of Sciences of the United States of AmericaJID - 7505876RN - 0 (DNA, Complementary)SB - IMMH - AnimalsMH - Base SequenceMH - Chiroptera/classification/*geneticsMH - DNA, Complementary

MH - EcosystemMH - *Evolution, MolecularMH - *FossilsMH - HumansMH - Molecular Sequence DataMH - PhylogenyPMC - PMC33452OID - NLM: PMC33452EDAT- 2001/05/17 10:00MHDA- 2001/07/20 10:01CRDT- 2001/05/17 10:00PHST- 2001/05/15 [aheadofprint]AID - 10.1073/pnas.111551998 [doi]AID - 111551998 [pii]PST - ppublishSO - Proc Natl Acad Sci U S A. 2001 May 22;98(11):6241-6. Epub 2001 May 15.