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FIGURE 12 in A new, narrowly endemic species of swamp-dwelling dusky salamander (Plethodontidae: Desmognathus) from the Gulf Coastal Plain of Mississippi and Alabama
FIGURE 12. Small specimen (MMNS 19992 /RAP1111) of Desmognathus valentinei from headwaters of West Creek, Harrison Co., Mississippi, showing typical juvenile coloration of "pale auriculatus" sensu Valentine (1963). Note strong overall similarity to D. conanti C (e.g., AMNH 193773/RAP1108; Fig. 11) on superficial examination.
FIGURE 4 in A new, narrowly endemic species of swamp-dwelling dusky salamander (Plethodontidae: Desmognathus) from the Gulf Coastal Plain of Mississippi and Alabama
FIGURE 4. The easternmost confirmed locality in the floodplain of Chickasaw Creek (AL: Mobile), in the Mobile River drainage. The ravine tributary stream originates in sight of I-65, and during our visit the upper sections were filled with trash discarded from the roadway. Nevertheless, an abundant salamander community persists where the intact hydrology supports sufficient microhabitat and vegetation. The site is heavily trafficked by humans for recreation and in close proximity (<5 km) to urban development, industrial activity, and interstate runoff.
FIGURE 1 in A new, narrowly endemic species of swamp-dwelling dusky salamander (Plethodontidae: Desmognathus) from the Gulf Coastal Plain of Mississippi and Alabama
FIGURE 1. Recent and historical populations previously or currently assigned to Desmognathus auriculatus (see Grobman 1950) based on verified localities from recent publications (Beamer and Lamb 2008; Means et al. 2017; Beamer and Lamb 2020; Pyron et al. 2022a,b) and museum records and new sites reported here. Specimens of unknown identity in northwestern Florida (FL) are representative of numerous extirpated historical populations in the region. These are thought to have been D. auriculatus A (Means et al. 2017; Pyron et al. 2022b) but have not been confirmed genetically and may be D. pascagoula or additional new species. All populations historically assigned to D. auriculatus from west of the Mississippi that have been tested genetically represent D. conanti E (Hibbitts et al. 2015) and are not shown. Other states of the southeastern United States: AR (Arkansas), LA (Louisiana), MS (Mississippi), TN (Tennessee), AL (Alabama), GA (Georgia), SC (South Carolina), and NC (North Carolina).
FIGURE 3. The Number 12 in A new, narrowly endemic species of swamp-dwelling dusky salamander (Plethodontidae: Desmognathus) from the Gulf Coastal Plain of Mississippi and Alabama
FIGURE 3. The Number 12 Swamp locality (Franklin Creek in Grand Bay, AL: Mobile), showing another view of the characteristic blackwater stream/cypress swamp habitat.
FIGURE 8 in A new, narrowly endemic species of swamp-dwelling dusky salamander (Plethodontidae: Desmognathus) from the Gulf Coastal Plain of Mississippi and Alabama
FIGURE 8. Results from the ML analysis of the ND2/tRNA/COI fragment for the focal clade, including Desmognathus auriculatus A–C, D. brimleyorum, D. conanti C, D. pascagoula, and D. valentinei. All available specimens are included for D. brimleyorum, D. pascagoula, and D. valentinei, but only a single exemplar for the other lineages. Dots at nodes indicate strong support (BS>70% & SHL>85%). Inset maps show all known localities for D. brimleyorum, D. conanti C, D. pascagoula, and D. valentinei verified by molecular sequence data (type locality and recent sites), along with the potential historical locality for D. pascagoula. Note that RAP1106 (in bold) is the holotype (USNM 596040).
FIGURE 11 in A new, narrowly endemic species of swamp-dwelling dusky salamander (Plethodontidae: Desmognathus) from the Gulf Coastal Plain of Mississippi and Alabama
FIGURE 11. Comparative specimen (AMNH 193773/RAP1108) of Desmognathus conanti C from Pigeon Creek, Clarke Co., Alabama, showing typical coloration of Spotted Dusky Salamanders from the Gulf Coastal Plain (see Valentine, 1963). These populations are often confused with D. valentinei (e.g., by Kozak et al., 2005; see Beamer and Lamb, 2008), and could be mistaken for D. pascagoula. Distinguishing characters include elongated "duck-bill" snout; indistinct postocular stripe; heavy, irregular, light-colored lateral and ventrolateral flecking and speckling; ventrolateral row of "portholes" on the trunk barely distinguishable from flecking; and more slender, less compressed tail with faint lateral row of "portholes" barely visible.
FIGURE 13 in A new, narrowly endemic species of swamp-dwelling dusky salamander (Plethodontidae: Desmognathus) from the Gulf Coastal Plain of Mississippi and Alabama
FIGURE 13. The holotype (USNM 596040/RAP1106) of Desmognathus pascagoula in life on isolated (a) and naturalistic (b) backgrounds. Note the relative lack of distinct "portholes" in the dorsolateral and lateral rows on the body and tail, and the bright orange postocular stripe. The ventrolateral row of "portholes" is partially visible on the body.
FIGURE 10. Individual boxplots showing the 9 in A new, narrowly endemic species of swamp-dwelling dusky salamander (Plethodontidae: Desmognathus) from the Gulf Coastal Plain of Mississippi and Alabama
FIGURE 10. Individual boxplots showing the 9 highest-loading (eigenvalues> 5) size-corrected linear measurements contributing to significant discrimination between all three related or sympatric species. While limited sample sizes preclude detailed significance tests or correction for multiple comparison, we include the notched boxplots here for a preliminary visual assessment of gross shape differences between the species relative to median values.
FIGURE 2 in A new, narrowly endemic species of swamp-dwelling dusky salamander (Plethodontidae: Desmognathus) from the Gulf Coastal Plain of Mississippi and Alabama
FIGURE 2. The original Grand Bay locality (Franklin Creek, AL: Mobile), showing the characteristic blackwater stream/ cypress swamp habitat, as distinct from the muddy, silted floodplain swamp at the type locality.
FIGURE 7 in A new, narrowly endemic species of swamp-dwelling dusky salamander (Plethodontidae: Desmognathus) from the Gulf Coastal Plain of Mississippi and Alabama
FIGURE 7. Results from conStruct analysis of 2,798 bi-allelic SNPs from 18 specimens. Overall layer contributions indicate K=2 significant genetic clusters, 10 Desmognathus valentinei from 6 populations, and 8 D. pascagoula from 5 populations.
FIGURE 9 in A new, narrowly endemic species of swamp-dwelling dusky salamander (Plethodontidae: Desmognathus) from the Gulf Coastal Plain of Mississippi and Alabama
FIGURE 9. Results from the LDA based on 17 linear morphometric measurements derived from Bingham et al. (2018). Significant separation (i.e., a majority of 95% ellipse volume) is observed between all three related or sympatric species, with the predominant loadings occurring primarily on trunk slenderness and craniofacial characteristics (see Fig. 10).
FIGURE 6 in A new, narrowly endemic species of swamp-dwelling dusky salamander (Plethodontidae: Desmognathus) from the Gulf Coastal Plain of Mississippi and Alabama
FIGURE 6. The northernmost locality, in the floodplain of Cowart Branch (MS: Greene) near the Leaf River. The site contains extensive bottomland blackwater cypress swamps.
Distribution. SE Kansas, SW & SE Missouri, E Oklahoma, Arkansas, E Texas, Louisiana, and SW Mississippi, USA, and extreme NE Tamaulipas, Mexico. in Cricetidae
Distribution. SE Kansas, SW & SE Missouri, E Oklahoma, Arkansas, E Texas, Louisiana, and SW Mississippi, USA, and extreme NE Tamaulipas, Mexico.
Subspecies and Distribution. O.p.palustrisHarlan,1837—widelydistributedinSIllinois,SKentucky,Tennessee,SNewJersey,SEPennsylvania,Delaware,EMaryland,EVirginia,ENorthCarolina,SouthCarolina,Georgia,Alabama,Mississippi,andNW&NFlorida,E&SEUSA. O.p.coloratusBangs,1898—SpeninsularandtheLowerKeysofFlorida,USA. O.p.natatorF.M.Chapman,1893—N&CpeninsularFlorida,USA. O.p.planirostrisHamilton,1955—limitedtoasmallareaonWCcoastofpeninsularFlorida,USA. O. p. sanibeli Hamilton, 1955 — Sanibel I on the SW coast of peninsular Florida, USA. in Cricetidae
Subspecies and Distribution. O.p.palustrisHarlan,1837—widelydistributedinSIllinois,SKentucky,Tennessee,SNewJersey,SEPennsylvania,Delaware,EMaryland,EVirginia,ENorthCarolina,SouthCarolina,Georgia,Alabama,Mississippi,andNW&NFlorida,E&SEUSA. O.p.coloratusBangs,1898—SpeninsularandtheLowerKeysofFlorida,USA. O.p.natatorF.M.Chapman,1893—N&CpeninsularFlorida,USA. O.p.planirostrisHamilton,1955—limitedtoasmallareaonWCcoastofpeninsularFlorida,USA. O. p. sanibeli Hamilton, 1955 — Sanibel I on the SW coast of peninsular Florida, USA.
Subspecies and Distribution. R.h.humulisAudubon&Bachman,1841—fromextremeNEArkansas,Mississippi,andSELouisianaEtoCVirginiaandStoSCFlorida(SEUSA). R.h.merriamiJ.A.Allen,1895—EOklahoma,ETexas,WArkansas,andW&SCLouisiana(SUSA). R. h. virginianus A. H. Howell, 1940 — SW Maryland and E Virginia (E USA). in Cricetidae
Subspecies and Distribution. R.h.humulisAudubon&Bachman,1841—fromextremeNEArkansas,Mississippi,andSELouisianaEtoCVirginiaandStoSCFlorida(SEUSA). R.h.merriamiJ.A.Allen,1895—EOklahoma,ETexas,WArkansas,andW&SCLouisiana(SUSA). R. h. virginianus A. H. Howell, 1940 — SW Maryland and E Virginia (E USA).
Subspecies and Distribution. O.n.nuttalliHarlan,1832—fromSVirginiaStoSWAlabama(SEUSA). O.n.aureolusAudubon&Bachman,1841—fromKentuckyandWestVirginiaStoNAlabamaandNGeorgia(EUSA). O.n.flammeusGoldman,1941—EOklahoma,SWMissouri,andArkansas(CUSA). O.n.flondanusPackard,1969—Florida(SEUSA). O. n. lisae Packard, 1969 — from SE Missouri and S Illinois S to NE Texas, Louisiana, and Mississippi (SC USA). in Cricetidae
Subspecies and Distribution. O.n.nuttalliHarlan,1832—fromSVirginiaStoSWAlabama(SEUSA). O.n.aureolusAudubon&Bachman,1841—fromKentuckyandWestVirginiaStoNAlabamaandNGeorgia(EUSA). O.n.flammeusGoldman,1941—EOklahoma,SWMissouri,andArkansas(CUSA). O.n.flondanusPackard,1969—Florida(SEUSA). O. n. lisae Packard, 1969 — from SE Missouri and S Illinois S to NE Texas, Louisiana, and Mississippi (SC USA).
Subspecies and Distribution. M.p.pinetorumLeConte,1830—SVirgin-1a,extremeETennessee,NorthCarolina,SouthCarolina,SEMississippi,SAlabama,andmostofGeorgia,USA. M.p.auricularisBailey,1898—SIndiana,SWOhio,extremeSEMissouri,mostofKentucky,E&SArkansas,mostofTennessee,ETexas,Louisiana,Mississippi,NAlabama,andNWGeorgia,USA. M.p.carbonariusHandley,1952—extremeSEOhio,extremeWWestVirginia,EKentucky,andETennessee,USA. M.p.nemoralisBailey,1898—extremeSEMinnesota,SWWisconsin,Iowa,SENebraska,EKansas,Missouri,EOklahoma,Arkansas,andNETexas,USA. M.p.parvulusA.H.Howell,1916—SGeorgiaandNFlorida,USA. M.p.scalopsoidesAudubon&Bachman,1841—extremeSOntarioandextremeSQuebec,Canada,andfromSWisconsinandIllinoisEtotheAtlanticcoastfromextremeSWMainetoSVirginia,USA. M. p. schmidti H. H. T. Jackson, 1941 — WC Wisconsin, USA. in Cricetidae
Subspecies and Distribution. M.p.pinetorumLeConte,1830—SVirgin-1a,extremeETennessee,NorthCarolina,SouthCarolina,SEMississippi,SAlabama,andmostofGeorgia,USA. M.p.auricularisBailey,1898—SIndiana,SWOhio,extremeSEMissouri,mostofKentucky,E&SArkansas,mostofTennessee,ETexas,Louisiana,Mississippi,NAlabama,andNWGeorgia,USA. M.p.carbonariusHandley,1952—extremeSEOhio,extremeWWestVirginia,EKentucky,andETennessee,USA. M.p.nemoralisBailey,1898—extremeSEMinnesota,SWWisconsin,Iowa,SENebraska,EKansas,Missouri,EOklahoma,Arkansas,andNETexas,USA. M.p.parvulusA.H.Howell,1916—SGeorgiaandNFlorida,USA. M.p.scalopsoidesAudubon&Bachman,1841—extremeSOntarioandextremeSQuebec,Canada,andfromSWisconsinandIllinoisEtotheAtlanticcoastfromextremeSWMainetoSVirginia,USA. M. p. schmidti H. H. T. Jackson, 1941 — WC Wisconsin, USA.
Subspecies and Distribution. O.z.zibethicusLinnaeus,1766—fromextremeSEManitoba,COntario,andCQuebec,Canada,SthroughEUSAtoextremeNETexas,extremeNLouisiana,Mississippi,Alabama,NGeorgia,andNWSouthCarolina. O.z.albusSabine,1823—Saskatchewan, Manitoba,andNOntario,Canada. O.z.aquiloniusBangs,1899—NQuebecandLabrador,Canada. O.z.bernard:Goldman,1932—SNevada,SECalifornia,andWArizona,USA,andNBajaCaliforniaandextremeNWSonora,Mexico. 0.z.aannamominusHollister,1910—fromSEAlberta,SSaskatchewan,andSManitoba,Canada,SthroughCUSAtoNTexas. O.z.goldmaniHuey,1938—SWUtah,SENevada,andNWArizona,USA. O.z.macrodonMerriam,1897—fromSEPennsylvaniaSalongthecoasttoNorthCarolina,USA. O.z.mergensHollister,1910—SEOregon,NC&NECalifornia,NNevada,andextremeNWUtah,USA. O.z.obscurusBangs,1894—Newfoundland,Canada. O.z.ocapitalisElliot,1903—WWashingtonandWOregon,USA. O.z.osoyoosensisLord,1863—fromSBritishColumbiaandSWAlberta,Canada,S alongtheRockyMtstoNNewMexico,USA. O.z.pallidusMearns,1890—fromSCArizonaEtoWNewMexico,USA. O.z.nipensisBailey,1902—fromCNewMexico,USAStoNChihuahua,Mexico. O.z.nwaliciusBangs,1895—fromSEcoastalTexasEtoextremeNWFlorida,USA. O.z.spatulatusOsgood,1900—fromPacificCoastofCAlaska,USA,SEtoSBritishColumbia,AlbertaandWSaskatchewan,Canada. O. z. zalophus Hollister, 1910 — SW & SC Alaska, USA. Introduced to most of Europe, Siberia, C Asia, Russian Far East, China, NE Korea, Japan (Honshu), S Chile, and S Argentina. in Cricetidae
Subspecies and Distribution. O.z.zibethicusLinnaeus,1766—fromextremeSEManitoba,COntario,andCQuebec,Canada,SthroughEUSAtoextremeNETexas,extremeNLouisiana,Mississippi,Alabama,NGeorgia,andNWSouthCarolina. O.z.albusSabine,1823—Saskatchewan, Manitoba,andNOntario,Canada. O.z.aquiloniusBangs,1899—NQuebecandLabrador,Canada. O.z.bernard:Goldman,1932—SNevada,SECalifornia,andWArizona,USA,andNBajaCaliforniaandextremeNWSonora,Mexico. 0.z.aannamominusHollister,1910—fromSEAlberta,SSaskatchewan,andSManitoba,Canada,SthroughCUSAtoNTexas. O.z.goldmaniHuey,1938—SWUtah,SENevada,andNWArizona,USA. O.z.macrodonMerriam,1897—fromSEPennsylvaniaSalongthecoasttoNorthCarolina,USA. O.z.mergensHollister,1910—SEOregon,NC&NECalifornia,NNevada,andextremeNWUtah,USA. O.z.obscurusBangs,1894—Newfoundland,Canada. O.z.ocapitalisElliot,1903—WWashingtonandWOregon,USA. O.z.osoyoosensisLord,1863—fromSBritishColumbiaandSWAlberta,Canada,S alongtheRockyMtstoNNewMexico,USA. O.z.pallidusMearns,1890—fromSCArizonaEtoWNewMexico,USA. O.z.nipensisBailey,1902—fromCNewMexico,USAStoNChihuahua,Mexico. O.z.nwaliciusBangs,1895—fromSEcoastalTexasEtoextremeNWFlorida,USA. O.z.spatulatusOsgood,1900—fromPacificCoastofCAlaska,USA,SEtoSBritishColumbia,AlbertaandWSaskatchewan,Canada. O. z. zalophus Hollister, 1910 — SW & SC Alaska, USA. Introduced to most of Europe, Siberia, C Asia, Russian Far East, China, NE Korea, Japan (Honshu), S Chile, and S Argentina.
Subspecies and Distribution. S.a.aquaticusLinnaeus,1758—Massachusetts,SENewYork,Connecticut,RhodeIsland,EPennsylvania,NewJersey,Maryland,Delaware,WestVirginia,Virginia,EKentucky,WNorthCarolina,ETennessee,NWSouthCarolina,andNGeorgia(USA). S.a.alleniR.H.Baker,1951—SCTexas(USA). S.a.anastasaeBangs,1898—AnastasiaI,Florida(USA). S.a.aereusBangs,1896—SKansas,Oklahoma,NETexas,Arkansas,andLouisiana(USA). S.a.australisF.M.Chapman,1893—SEGeorgiaandmostofFlorida(USA). S.a.bassiA.H.Howell,1939—knownonlyfromthetypelocalityinS.a.County,EFlorida(USA). S.a.carytH.H.T.Jackson,1914—SSouthDakota,SEWyoming,NEColorado,W&CNebraska,andNWKansas(USA). S.a.howelliH.H.T.Jackson,1914—Mississippi,SELouisiana,Alabama,extremeNWFlorida,W&NGeorgia,mostofNorthandSouthCarolina,andextremeSCVirginia(USA). S.a.inflatusH.H.T.Jackson,1914—extremeSTexas(USA)andNETamaulipas(Mexico). S.a.machrinoidesH.H.T.Jackson,1914—SMinnesota,ENebraska,Iowa,EKansas,Missouri,NEOklahoma,andNArkansas(USA). S.a.machrinusRafinesque,1832—SOntario(Canada)andWisconsin,Michigan,IIlinois,Indiana,Ohio,W&CKentucky,WWestVirginia,andW&CTennessee(USA). S.a.montanusR.H.Baker,1951—NCoahuila(Mexico)andadjacentTexas(USA). S.a.nanusW.B.Davis,1942—ETexas(USA). S.a.parvusRhoads,1894—EFlorida(USA). S.a.porteriSchwartz,1952—SFlorida(USA). S. a. texanusJ. A. Allen, 1893 — Presidio County, SW Texas (USA). in Talpidae
Subspecies and Distribution. S.a.aquaticusLinnaeus,1758—Massachusetts,SENewYork,Connecticut,RhodeIsland,EPennsylvania,NewJersey,Maryland,Delaware,WestVirginia,Virginia,EKentucky,WNorthCarolina,ETennessee,NWSouthCarolina,andNGeorgia(USA). S.a.alleniR.H.Baker,1951—SCTexas(USA). S.a.anastasaeBangs,1898—AnastasiaI,Florida(USA). S.a.aereusBangs,1896—SKansas,Oklahoma,NETexas,Arkansas,andLouisiana(USA). S.a.australisF.M.Chapman,1893—SEGeorgiaandmostofFlorida(USA). S.a.bassiA.H.Howell,1939—knownonlyfromthetypelocalityinS.a.County,EFlorida(USA). S.a.carytH.H.T.Jackson,1914—SSouthDakota,SEWyoming,NEColorado,W&CNebraska,andNWKansas(USA). S.a.howelliH.H.T.Jackson,1914—Mississippi,SELouisiana,Alabama,extremeNWFlorida,W&NGeorgia,mostofNorthandSouthCarolina,andextremeSCVirginia(USA). S.a.inflatusH.H.T.Jackson,1914—extremeSTexas(USA)andNETamaulipas(Mexico). S.a.machrinoidesH.H.T.Jackson,1914—SMinnesota,ENebraska,Iowa,EKansas,Missouri,NEOklahoma,andNArkansas(USA). S.a.machrinusRafinesque,1832—SOntario(Canada)andWisconsin,Michigan,IIlinois,Indiana,Ohio,W&CKentucky,WWestVirginia,andW&CTennessee(USA). S.a.montanusR.H.Baker,1951—NCoahuila(Mexico)andadjacentTexas(USA). S.a.nanusW.B.Davis,1942—ETexas(USA). S.a.parvusRhoads,1894—EFlorida(USA). S.a.porteriSchwartz,1952—SFlorida(USA). S. a. texanusJ. A. Allen, 1893 — Presidio County, SW Texas (USA).
Subspecies and Distribution. B.c.carolinensisBachman,1837—SEUSA,EoftheMississippiRiver(SIllinois,WKentucky,WTennessee,Mississippi,SELouisiana,SAlabama,SGeorgia,NFlorida,SouthCarolina,ENorthCarolina,andSEVirginia). B. c. minima Lowery, 1943 — SE USA, W of the Mississippi River (SE Missouri, Arkansas, most of Louisiana, and E Texas). in Soricidae
Subspecies and Distribution. B.c.carolinensisBachman,1837—SEUSA,EoftheMississippiRiver(SIllinois,WKentucky,WTennessee,Mississippi,SELouisiana,SAlabama,SGeorgia,NFlorida,SouthCarolina,ENorthCarolina,andSEVirginia). B. c. minima Lowery, 1943 — SE USA, W of the Mississippi River (SE Missouri, Arkansas, most of Louisiana, and E Texas).
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Allen Brain Atlas
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