Gayana 73(2): 180 - 199, 2009 ISSN 0717-652X
DIVERSITY AND DISTRIBUTION OF THE MOUSE OPOSSUMS OF THE GENUS THYLAMYS (DIDELPHIMORPHIA, DIDELPHIDAE) IN NORTH- EASTERN AND CENTRAL ARGENTINA
DIVERSIDAD Y DISTRIBUCION DE LAS MARMOSAS DEL GENERO THYLAMYS (DIDELPHIMORPHIA, DIDELPHIDAE) EN EL NORESTE Y CENTRO DE ARGENTINA
Pablo Teta1**XLOOHUPR'¶(OtD2, David Flores1 1RpGH/D6DQFKD3
1 Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Avenida Angel Gallardo 470 (C1405DJR) Buenos $LUHV$UJHQWLQD 2 'HSDUWDPHQWRGH=RRORJtD8QLYHUVLGDGGH&RQFHSFLyQFDVLOOD&&RQFHSFLyQ&KLOH 3 'HSDUWPHQWRI%LRORJLFDO6FLHQFHV7H[DV7HFK8QLYHUVLW\32%R[/XEERFN7H[DV86$ E-mail: DQWKHFD#\DKRRFRPDr
ABSTRACT
Phylogenetic analysis of a fragment of the mitochondrial genome and qualitative and quantitative assessments of morphological variation suggest that, in its current conception, Thylamys pusillus 'HVPDUHVW LVDFRPSOH[RIDW OHDVWWKUHHVSHFLHV,QWKHWD[RQRPLFDUUDQJHPHQWSURSRVHGLQWKLVZRUNWKHSRSXODWLRQVLQWKH$UJHQWLQHDQSURYLQFHV of Entre Ríos and Corrientes are here referred to T citellus (Thomas, 1912), while the small Thylamys that lives in the Argentinean Dry Chaco are provisionally referred to Tpulchellus &DEUHUD ,QRXUVFKHPHThylamys pusillus is UHVWULFWHGWRWKH%ROLYLDQDQG3DUDJXD\DQ&KDFRDQGWKHYLFLQLWLHVRIQRUWKHUQ)RUPRVDSURYLQFHLQ$UJHQWLQD:HSURYLGH emended diagnosis for T citellus and Tpulchellus, together with detailed morphological descriptions and discuss their distinctiveness from other species of Thylamys,QDGGLWLRQZHLQFOXGHGQHZGLVWULEXWLRQDOGDWD
KEYWORDS $UJHQWLQDPRXVHRSRVVXPVSHFLHVOLPLWVWD[RQRP\
RESUMEN
(ODQiOLVLV¿ORJHQpWLFRGHXQIUDJPHQWRGHOJHQRPD\HOHVWXGLRFXDOLWDWLYR\FXDQWLWDWLYRGHODPRUIRORJtDH[WHUQD\ craneana sugiere que, en su actual acepción, Thylamys pusillus (Desmarest, 1804) es un complejo de por lo menos tres HVSHFLHV(QHODUUHJORWD[RQyPLFRSURSXHVWRHQHVWHWUDEDMRODVSREODFLRQHVHQODVSURYLQFLDVDUJHQWLQDVGH(QWUH5tRV y Corrientes son referidas como T citellus (Thomas, 1912), mientras que una forma pequeña de Thylamys que habita en el Chaco Seco de Argentina es provisoriamente referida como Tpulchellus &DEUHUD (QHVWHHVTXHPDOD distribución de Thylamys pusillus es restringida al Chaco de Bolivia, Paraguay y áreas adyacentes de Argentina en el QRUGHVWHGHODSURYLQFLDGH)RUPRVD6HSURYHHQGLDJQRVLVHQPHQGDGDVSDUDT citellus y Tpulchellus, conjuntamente con una detallada descripción morfológica de ambas entidades y una discusión de las diferencias con otras especies de Thylamys3RUXOWLPRVHLQFOX\HQQXHYRVGDWRVGHGLVWULEXFLyQ
PALABRAS CLAVE:$UJHQWLQDPDUPRVDVOtPLWHGHHVSHFLHVWD[RQRPtD
180 7D[RQRP\RIThylamys: TETA,P.ET AL.
INTRODUCTION al. 2005a, Flores et al.&UHLJKWRQ *DUGQHU 2008) restrict the name pusillus to the central and The genus Thylamys Gray 1843 comprises small northeastern populations from Argentina, western mouse opossums with some distinctive traits, 3DUDJXD\DQGHDVWHUQ%ROLYLD8QGHUWKLVFRQFHSWLRQ including a characteristic tricolor fur pattern, T. pusillus includes, as subjective junior synonyms, capacity to store fat in the tail, uniformly narrow WKHQRPLQDOWD[DMarmosa citella 7KRPDV nasals with subparallel lateral margins, large Marmosa janetta pulchella &DEUHUD DQG SRVWHURODWHUDOSDODWDOIRUDPLQDH[WHQGLQJDQWHULRUO\ Marmosa verax7KRPDV VHH&UHLJKWRQ between M4 protocones, and other morphological *DUGQHU 3DUWRIWKHFRQIXVLRQUHJDUGLQJ characters in body proportions, skull, dentition, and the biological and geographic limits of T. pusillus SRVWFUDQLXP &UHLJWKRQ *UDGQHU)ORUHV GHULYHVIURPWKHDEVHQFHRIDW\SHVSHFLPHQ,Q 9RVV -DQVD 8QOLNHPRVWRWKHU6RXWK fact, the name pusillus was proposed on the basis of American mouse opossums, which are found in $]DUD¶VGHVFULSWLRQRIWKH³PLFRXUpQDLQ´RU³HQDQR´ tropical or subtropical moist forests, Thylamys (Azara, 1801, 1802), without the formal designation principally inhabits semi-arid and open areas and RIDW\SHVSHFLPHQ*LYHQWKLV9RVVet al. (in press) occurs at temperate latitudes (Flores et al. 2007, designed a neotype for pusillus from Trans-Chaco &UHLJKWRQ *DUGQHU +LJKZD\NP1:RI9LOOD+D\HV 'HSDUWDPHQWR ThylamysKDVEHHQXQGHUWD[RQRPLFUHYLVLRQ %RTXHUyQ3DUDJXD\ ,QWKHVDPHFRQWULEXWLRQWKH since Tate (1933) recognized the elegans species authors suggest that some Argentinean populations group as part of the genus Marmosa6XEVHTXHQW of pusillus-like Thylamys may represent a different systematic works, based on morphological, species than the nominotypical form that occurs VHURORJLFDODQGFKURPRVRPDOHYLGHQFHFRQ¿UPHG LQ3DUDJXD\DQGHDVWHUQ%ROLYLD2QWKHRWKHU the distinctiveness of Thylamys within the didelphid hand, Argentinean populations from Entre Ríos radiation (Reig et al. *DUGQHU &UHLJKWRQ and Corrientes provinces (Mesopotamia), usually 1989, Palma et al. 3K\ORJHQHWLFVWXGLHV synonymyzed in the literature with pusillus, have EDVHGRQJHQHWLFGDWDFRQ¿UPHGWKHPRQRSK\O\ DORQJDQGFRPSOH[WD[RQRPLFKLVWRU\EHLQJ of the genus, and elucidated some basic questions alternatively included in Marmosa marmota, regarding relationships among its species and their Thylamys pusillus, or considered a separated species OLPLWV .LUVFK 3DOPD -DQVD 9RVV under the name of M. citella 7KRPDV 3DOPD
181 Gayana 73(2), 2009
FIGURE 1. Recording localities for the specimens of Thylamys citellus (circles), T. pulchellus (triangles), and T. pusillus VTXDUHV UHYLVHGLQWKLVZRUN QXPEHUVFRUUHVSRQGVWRWKRVHRIWKH$SSHQGL[ DQGDGGLWLRQDOORFDOLWLHVIRUThylamys pulchellus (from Braun et al. $UJHQWLQD &DWDPDUFD &DSD\iQ&KXPELFKDNP(RI+LJKZD\DORQJ 6: 6DQ-XDQ &DXFHWH4XHEUDGDGH/DV)ORUHVNP(DQGNP1*XD\DPDV FD 6 : 6DQWLDJRGHO(VWHUR $OEHUGLNP6NP(3DPSDGH/RV*XDQDFRV 6: &KR\D6DOLQDVGH$PEDUJDVWDDSSNP6(&HUUR5LFR 6: DQG Tpusillus 3DUDJXD\ %RTXHUyQ 7UDQV&KDFR+LJKZD\NP1:9LOOD+D\HV 6 : W\SHORFDOLW\¿[HGE\QHRW\SHVHOHFWLRQ>9RVVet al.LQSUHVV@
FIGURA 1/RFDOLGDGHVGHUHJLVWURSDUDORVHMHPSODUHVGHThylamys citellus (círculos), T. pulchellus (triángulos) y T. pusillus FXDGUDGRV UHYLVDGRVHQHVWHWUDEDMR ORVQ~PHURVFRUUHVSRQGHQDORVDQRWDGRVHQHO$SpQGLFH \ORFDOLGDGHV adicionales para Thylamys pulchellus (tomadas de Braun et al. $UJHQWLQD &DWDPDUFD &DSD\iQ&KXPELFKD NP(RI+LJKZD\DORQJ 6: 6DQ-XDQ &DXFHWH4XHEUDGDGH/DV)ORUHV NP(DQGNP1*XD\DPDV FD6 : 6DQWLDJRGHO(VWHUR $OEHUGLNP6NP( 3DPSDGH/RV*XDQDFRV 6: &KR\D6DOLQDVGH$PEDUJDVWDDSSNP6(&HUUR5LFR 6: \Tpusillus 3DUDJXD\ %RTXHUyQ 7UDQV&KDFR+LJKZD\NP1:9LOOD +D\HV : ORFDOLGDGWLSR¿MDGDSRUVHOHFFLyQGHQHRWLSR>9RVVet al.HQSUHQVD@ of Thylamys pallidior VHH$SSHQGL[ DQGXVHG -DQVD 6WDQGDUGH[WHUQDOPHDVXUHVIRU diagnostic traits annotated in the literature for T mammals were recorded from field catalogs or macrurus (Voss et alLQSUHVV VSHFLPHQWDJV +% KHDGDQGERG\OHQJWK 7/ WDLO length, HF: hind foot length (including the claw), and MORPHOLOGIC DATA ANALYSIS1RPHQFODWXUHXVHGWR (/ HDUOHQJWK(LJKWVNXOODQGGHQWDOPHDVXUHPHQWV describe the skull and its structures follows Voss were recorded following Voss et al. D WKHVH
182 7D[RQRP\RIThylamys: TETA,P.ET AL.
TABLE I.'HVFULSWLYHVWDWLVWLFVIRUH[WHUQDODQGFUDQLDOYDULDEOHVRIDGXOWPDOHVDQGIHPDOHV DJHFODVVHVDQG RI Thylamys citellus, T. pulchellus, and T. pusillus6HHPDWHULDOVDQGPHWKRGVIRUWKHH[SODQDWLRQRIWKHDEEUHYLDWXUHV 6G VWDQGDUGGHYLDWLRQ
TABLA I.(VWDGtVWLFRVGHVFULSWLYRVSDUDYDULDEOHVH[WHUQDV\FUDQHDQDVGHHMHPSODUHVDGXOWRVPDFKRV\KHPEUDV FODVHV de edad 6 y 7) de Thylamys citellus, Tpulchellus y Tpusillus9pDVHPDWHULDOHV\PpWRGRVSDUDXQDH[SOLFDFLyQGHODV DEUHYLDWXUDV 6G GHVYLDFLyQHVWiQGDU
MI- HB T HF EL CBL ZB LIB PL PB MTR LM M3 Thylamys Mean citellus Minimum 0D[LPXP Sd N 555555555555 Thylamys Mean pulchellus Minimum 0D[LPXP Sd N 1111911999999109 Thylamys Mean pusillus Minimum 0D[LPXP 6,5 Sd N 383838365759616262624862 are: condylobasal length (CBL), palatal length (PL), species of ThylamysZHUHDQDO\]HG 7DEOH,, 2XU least interorbital breadth (LIB), palatal breadth (PB), sampling lacks sequences representing T. macrurus, ]\JRPDWLFEUHDGWK =% PD[LOODU\WRRWKURZOHQJWK T. sponsorius, T. tatei, and T. velutinus+DSORW\SHV (MTR), length of molars (LM), and length of M1-M3 recovered from 10 specimens representing 10 other 00 ([WHUQDODQGFUDQLRGHQWDOPHDVXUHPHQWV GLGHOSKLGJHQHUDZHUHXVHGDVRXWJURXSV7KLUW\ DUHVXPPDUL]HGLQ7DEOH,$SULQFLSDOFRPSRQHQW three of the 37 sequences analyzed were retrieved analysis was conducted using the software Statistica IURP*HQEDQN 7DEOH, :HJHQHUDWHGWKHVHTXHQFHV (StatSoft 2001), performed on the eight craniodental of the following three specimens from the Argentine PHDVXUHPHQWVSUHYLRXVO\GH¿QHGDQGFRQVLGHULQJ Mesopotamia currently assigned to T. pusillus: only adult specimens (ages 6-7, sensu Tribe, 1990) CNP 1920, MACN 23459, and MACN 23460 as ZLWKRXWPLVVLQJYDOXHV3ULQFLSDOFRPSRQHQWV well as one of Cryptonanus chacoensis (GD 521) ZHUHH[WUDFWHGIURPDYDULDQFHFRYDULDQFHPDWUL[ XVHGDVSDUWRIWKHRXWJURXS/DERUDWRU\SURFHGXUHV and computed by using the craniodental variables IROORZHGWKRVHRI'¶(OtD 3DUGLxDV DIWHUWUDQVIRUPDWLRQWRWKHLUQDWXUDOORJDULWKPV New sequences were translated to amino-acids 1RPHQFODWXUHRIFRORUVIROORZV5LGJZD\ DQGQRHYLGHQFHRIQRQIXQFWLRQDOLW\ZDVIRXQG PHYLOGENETIC ANALYSES. Phylogenetic analyses were Sequences were aligned using Clustal X (Thompson EDVHGRQWKH¿UVWEDVHSDLUVRIWKHPLWRFKRQGULDO et al. 1997) with default values for all alignment JHQHWKDWFRGHVIRUF\WRFKURPHE6HTXHQFHVRI SDUDPHWHUV QRDGMXVWPHQWE\H\HZDVQHHGHG VSHFLPHQVEHORQJLQJWRVL[FXUUHQWO\UHFRJQL]HG Observed percentage of sequence divergence was
183 Gayana 73(2), 2009 calculated with MEGA 4 (Tamura et al. 2007) western of Paraná River (southern Chaco) with the LJQRULQJWKRVHVLWHVZLWKPLVVLQJGDWD6HTXHQFH H[FHSWLRQRI¿YHVSHFLPHQVIURP)RUPRVD3URYLQFH DOLJQPHQWZDVVXEMHFWHGWRPD[LPXPSDUVLPRQ\ near the Argentinean-Paraguayan border, which 03 )DUULV DQG%D\HVLDQ +XHOVHQEHFNet groups with the Paraguayan samples to form the al. DQDO\VHV&KDUDFWHUVXVHGLQ03DQDO\VLV WKLUGJURXSIURPWKHQRUWKHUQ&KDFR,QWKHELYDULDWH ZHUHWUHDWHGDVXQRUGHUHGDQGHTXDOO\ZHLJKWHG03 SORWRIFRPSRQHQWV,DQG,, )LJ WKHRYHUODSLQJ was conducted in PAUP* (Swofford 2000) with 200 is scarce, and the groups are mostly separated on the replicates of heuristic search with random addition ¿UVWFRPSRQHQW+HUHWKHPRVWGLVVLPLODUJURXSLVWKH RIVHTXHQFHVDQG7%5EUDQFKVZDSSLQJ1RGDO one including specimens from Corrientes and Entre support was assessed with 1000 pseudoreplicates 5tRV2QWKHRWKHUKDQGWKHVRXWKHUQDQGQRUWKHUQ RI-DFNNQLIH -. )DUULVet al. 1996), each with 33 Chacoan groups are poorly differentiated along the % of data deletion and three replicates of random ¿UVWDQGVHFRQGFRPSRQHQWVDVZHOODVDORQJWKH DGGLWLRQRIVHTXHQFHVDQG7%5EUDQFKVZDSSLQJ ¿UVWDQGWKLUGFRPSRQHQWV )LJ +RZHYHUWKH Bayesian analysis was conducted with MrBayes 3 southern group in always placed on the rigth part of 5RQTXLVW +XHOVHQEHFN E\PHDQVRIWZR WKHSORW%DVLFDOO\WKHWKUHHJURXSVGLVFULPLQDWHE\ independent runs with three heated and one cold size in several cranial measurements as CBL, PL and 0DUNRYFKDLQVHDFK$PRGHOZLWKVL[FDWHJRULHV =% 7DEOH,,, of base substitution, a gamma-distributed rate parameter, and a proportion of invariant sites was PHYLOGENETIC INFERENCE0D[LPXPSDUVLPRQ\ VSHFL¿HG DOOPRGHOSDUDPHWHUVZHUHHVWLPDWHGLQ DQDO\VLVUHFRYHUHGVKRUWHVWWUHHV OHQJWK 0U%D\HV8QLIRUPLQWHUYDOSULRUVZHUHDVVXPHG VWHSV &, 5, 7KHVWULFWFRQVHQVXV IRUDOOSDUDPHWHUVH[FHSWEDVHFRPSRVLWLRQDQG RIWKHVHWUHHV )LJD VKRZVDZHOOVXSSRUWHG GTR parameters, which assumed a Dirichlet -. Thylamys clade, which has 19 nodes of SURFHVVSULRU&KDLQVZHUHDOORZHGWRUXQIRUWKUHH ZKLFK¿YHLQYROYHSRO\WRPLHV6SHFLPHQVFXUUHQWO\ PLOOLRQJHQHUDWLRQV WUHHVZHUHVDPSOHGHYHU\ assigned to T pusillus form a strongly supported JHQHUDWLRQVIRUHDFKFKDLQ7RFKHFNWKDWHDFKUXQ -. FODGHVLVWHUWRDFODGH -. IRUPE\ converged on a stable log-likelihood value, we the other two southern cone species included in the plotted the log-likelihood values against generation analysis, Tpallidior and Telegans5HODWLRQVKLSV WLPHIRUHDFK7KHILUVWRIWKHWUHHVZHUH RIWKHRWKHUVSHFLHVDUHXQUHVROYHG:LWKLQWKH discarded as burn-in and the remaining trees were clade formed by specimens currently assigned to T used to compute a 50% majority rule consensus tree pusillus there are three main clades geographically and obtain posterior probability (PP) estimates for allopatric and coincident with those found in the HDFKFODGH PRUSKRPHWULFDQDO\VLV6SHFLPHQVIURP$UJHQWLQD ZHVWRI3DUDQi5LYHU LHIURP&DWDPDUFD Santiago del Estero, and San Juan Provinces) form RESULTS DVWURQJO\VXSSRUWHGFODGH -. 6SHFLPHQV IURP$UJHQWLQH0HVRSRWDPLD LHHDVWRI3DUDQi MORPHOMETRIC ANALYSIS7KHHLJHQYDOXHVRIWKH¿UVW River in Entre Ríos province) form another strongly WKUHHSULQFLSDOFRPSRQHQWVZHUH VXSSRUWHGFODGH -. %RWKWKHZHVWHUQ3DUDQi DQG UHVSHFWLYHO\ and Mesopotamian clades are sister groups in a 7RJHWKHUWKHVHWKUHHD[HVDFFRXQWIRU RIWKH UHODWLYHO\ZHOOVXSSRUWHGFODGH -. ZKLFKLV total variance in our log-transformed craniodental sister to the clade formed by the single Paraguayan PHDVXUHPHQWGDWD/RDGLQJVRIWKHHLJKWFKDUDFWHUV VSHFLPHQLQFOXGHGLQWKHDQDO\VLV5HVXOWVRIWKH DQDO\]HGDORQJHDFKRIWKH¿UVWWKUHHFRPSRQHQWV %D\HVLDQDQDO\VLV )LJE DUHFRQJUXHQWZLWKWKRVH DUHVKRZQLQ7DEOH,,,$OWKRXJKWKHUHLVDQ RIWKH03DQDO\VLV WKHPDLQGLIIHUHQFHLVWKDWDVLV evident overlaping among the specimens included usually the case, the Bayesian tree is more resolved in the multivariate analysis, specimens referable LHT venustus appears as sister to T cinderella to Thylamys pusillus (as that species is currently T karimii is sister to the clade formed by the other understood) split in three main groups geographically Thylamys WKDQWKH03WUHH7KH%D\HVLDQDQDO\VLV allopatric: one group is formed by Mesopotamian is totally congruent with the MP tree in the topology specimens, another by all Argentinean samples obtained from the haplotypes of specimens currently
184 7D[RQRP\RIThylamys: TETA,P.ET AL.
TABLE II./LVWRIVSHFLPHQVIURPZKLFKKDSORW\SHVLQFOXGHGLQWKHSK\ORJHQHWLFDQDO\VHVZHUHUHFRYHUHG)RUHDFK VHTXHQFHLWV*HQEDQNDFFHVVLRQQXPEHUVDQGWKHLQGLFDWLRQRIVWXG\LQZKLFKLWZDVJDWKHUHGDUHSURYLGHG)RUGDWD of the specimens of ThylamysVHTXHQFHGLQWKLVVWXG\VHHWKH$SSHQGL[7KHVSHFLPHQRICryptonanus chaoensis (GD *' FDWDORJQXPEHURI*XLOOHUPR'¶(OtD ZDVFROOHFWHGLQ3DUDJXD\&DD]DSi(VWDQFLD'RV0DUtDVNP6 GHODVFDVDVFRVWDGHO5tR7HELFXDU\ 6:
TABLA II./LVWDGHHVSHFtPHQHVGHORVFXDOHVVHUHFREUDURQORVKDSORWLSRVLQFOXLGRVHQORVDQiOLVLV¿ORJHQpWLFRV3DUD WRGDVODVVHFXHQFLDVVHSURYHHQHOQ~PHURGHDFFHVRD*HQEDQN\VHLQGLFDHOHVWXGLRHQTXHIXHJHQHUDGD(QUHODFLyQD los datos de los especímenes de ThylamysVHFXHQFLDGRVHQHVWHHVWXGLRYHUHO$SpQGLFH(OHVSpFLPHQGHCryptonanus chaoensis *' *' Q~PHURGHFDWiORJRGH*XLOOHUPR'¶(OtD IXHFROHFWDGRHQ3DUDJXD\&DD]DSi(VWDQFLD'RV 0DUtDVNP6GHODVFDVDVFRVWDGHO5tR7HELFXDU\ 6:
7D[RQ Catalog number Accession Number Reference Ingroup 1 T. cinderella Arg 2234 AY803332 Braun et al. (2005a) 2 T. cinderella OMNH 22965 AY803325 Braun et al. (2005a) 3 T. cinderella OMNH 29975 AY803327 Braun et al. (2005a) 4 T. cinderella OMNH 29977 AY803329 Braun et al. (2005a) 5 T. citellus MACN 23459 GQ911594 This study 6 T. citellus MACN 23460 GQ911593 This study 7 T. citellus CNP 1920 GQ911595 This study 8 T. elegans MUSM 10738 AF434179 Meynard et al. (2002)
9 T. elegans 1. AF431929 Palma et al. (2002) 10 T. elegans 1. AF434176 Meynard et al. (2002) Carvalho et al. 11 T. karimii - EF051700 (unpublished) Carvalho et al. 12 T. karimii MN 36405 EF 114742 (unpublished) 13 T. pallidior CML 3192 AY803311 Braun et al. (2005a) 14 T. pallidior EP 440 AF431923 Palma et al. (2002) 15 T. pallidior FMNH 162495 AJ628368 Steiner et al. (2005)
16 T. pallidior 1. AF431924 Palma et al. (2002) 17 T. pallidior 1. AF431930 Palma et al. (2002) 18 T. pallidior OMNH 29957 AY803300 Braun et al. (2005) 19 T. pallidior OMNH 29964 AY803297 Braun et al. (2005) 20 T. pulchellus Arg 5420 AY803322 Braun et al. (2005) 21 T. pulchellus CML 3198 AY803323 Braun et al. (2005) 22 T. pulchellus OMNH 23479 AY803321 Braun et al. (2005) 23 T. pulchellus OMNH 23483 AY803319 Braun et al. (2005) 24 T. pulchellus OMNH 29961 AY803320 Braun et al. (2005)
185 Gayana 73(2), 2009
Continuación TABLA II
25 T. pusillus 7. AY803324 Braun et al. (2005) 26 T. venustus 1. AF431922 Palma et al. (2002) 27 T. venustus OMNH 29952 AY803334 Braun et al. (2005)
Outgroup 28 Chironectes minimus ROM FN-31677 AJ628363 Steiner et al. (2005) Cryptonanus chacoen- 29 GD 521 GQ911596 This study sis Gracilinanus micro- 30 MVZ 182055 AJ628369 Steiner et al. (2005) tarsus Lutreolina crassicau- 31 UMMZ 134018 AJ628364 Steiner et al. (2005) data 32 Marmosa lepida AMNH 273186 AJ606452 Steiner et al. (2005) Marmosa (Micoureus) 33 MNFS 181 U34673 Patton et al. (1996) demerarae 34 Marmosops impavidus MVZ 171408 U34669 Patton et al. (1996) Metachirus nudicau- 35 MNFS 40 U34672 Patton et al. (1996) datus 36 Monodelphis adusta MVZ 171412 U34677 Patton et al. (1996) 37 Philander opossum MNFS 146 U34679 Patton et al. (1996)
assigned to T. pusillus, although is worth noting KDVLWVW\SHORFDOLW\LQWKHSURYLQFHRI&RUULHQWHV that the Mesopotamian clade is only moderately The known distribution of T. citellus is restricted VXSSRUWHG 33 E\WKH3DUDQiDQG8UXJXD\ULYHUV2QWKHRWKHU hand, the small specimens coming from the western TAXONOMY7DNHQDVDZKROHRXUUHVXOWVDOORZXVWR VLGHRIWKH3DUDQi5LYHULQWKHPRUHH[WHQVLYH GLVWLQJXLVKWKUHHJURXSVZLWKLQWKHWD[RQFXUUHQWO\ Dry Chaco eco-region, are clearly distinguishable recognized as Thylamys pusillus7KLVVFHQDULRLV from Mesopotamian populations (T. citellus), and also supported by the fact that two of these groups from those in northern Formosa and Paraguay (T. GRQRW¿WZLWKWKHHPHQGHGGLDJQRVLVRITpusillus pusillus *HQHWLFGLYHUJHQFHDQGPRUSKRORJLFDO provided by Voss et al. LQSUHVV ,QDGGLWLRQWKH characters support the idea that such specimens specimens coming from the northern portion of the could be assigned to a species inhabiting most of $UJHQWLQH+XPLG&KDFRLQ)RUPRVD3URYLQFH )LJ the Dry Chaco eco-region in northern and central $SSHQGL[ DUHPRUSKRORJLFDOO\FORVHUWRT $UJHQWLQD7KHVHSRSXODWLRQVFDQEHDVVLJQHGWR pusillus IURP%ROLYLDDQG3DUDJXD\7KHUHPDLQLQJ T. pulchellus Cabrera, 1934, based on a specimen specimens, which are split in two discrete groups, from the dry Chacoan forest of Santiago del Estero ZRXOGFRUUHVSRQGWRWZRDGGLWLRQDOVSHFLHV7KH 3URYLQFH )LJ ZKRVHPRUSKRORJ\UHVHPEOHVWKDW genetic evidence and the morphological pattern RIH[DPLQHGYRXFKHUVVSHFLPHQVIURPVRXWKZHVWHUQ (see below), indicate that the populations of the Chaco, southeastern Salta, and Santiago del Estero Argentine Mesopotamia (Entre Rios and Corrientes VHH$SSHQGL[ ,QVXPPDU\ZLWKWKHHYLGHQFH 3URYLQFHV)LJ RQWKHHDVWHUQVLGHRI3DUDQi at hand, we recognize three distinct species within River, can be assigned to Tcitellus (Thomas, 1912), the Thylamys pusillus FRPSOH[ T citellus, T which is among the largest species of Thylamys and pulchellus, and T pusillus (sensu stricto %HORZ
186 7D[RQRP\RIThylamys: TETA,P.ET AL.
TABLE III. Results of principal components analysis of eight craniodental measurements of adult specimens of Thylamys DJHFODVHV Q 6HH0DWHULDOVDQG0HWKRGVIRUDEEUHYLDWLRQV
TABLA III. Resultados del análisis de componentes principales para ocho medidas craneodentarias de ejemplares adultos de Thylamys FODVHVGHHGDG Q 9pDVH0DWHULDOHV\0pWRGRVSDUDXQDH[SOLFDFLyQGHODVDEUHYLDWXUDV
PC I PC II PC III ZB -0,875088 0,230073 0,102571 LIB -0,714867 0,029994 0,467879 CBL -0,939601 0,201280 0,020560 PL -0,942238 0,139973 -0,007780 PB -0,782272 0,217435 -0,139311 MTR -0,858232 -0,288868 -0,058945 LM -0,290703 -0,865606 0,316361 M1-3 -0,566221 -0,378527 -0,651012 Eigenvalue 4,801102 1,137217 0,776697 % Variance 60,01378 14,21522 9,70872
FIGURE 2. Specimen scores of adult individuals of Thylamys DJHFODVHV Q IRUSULQFLSDOFRPSRQHQWVDQG DQGDQGH[WUDFWHGIURPWKHYDULDQFHFRYDULDQFHPDWUL[RIFUDQLRGHQWDOGLVWDQFHV VHHWH[WDQG7DEOH,,
FIGURA 2. Proyección de los ejes 1 y 2 y 1 y 3 del análisis de componentes principales generado a partir de una matriz de varianza-covarianza de 8 medidas craneo dentarias para ejemplares adultos de Thylamys FODVHVGHHGDG Q we offer emended diagnoses and comparisons for T $OW¶ )LJ citellus and Tpulchellus)RUWKHHPHQGHGGLDJQRVLV of T. pusillus see Voss et al. LQSUHVV TYPE LOCALITY. Goya, Corrientes Province, Argentina 6 : )LJ Thylamys citellus (Thomas, 1912) TAXONOMIC REMARKS. 7KH¿UVWThylamys specimen )LJV7DEOH,,,9 IURP&RUULHQWHV3URYLQFH$UJHQWLQDZDVLGHQWL¿HG by Thomas (1894) as Micoureus griseus Desmarest, HOLOTYPE.%0 FROOHFWHGth July, 7ZR\HDUVODWHU7KRPDV XVHGWKH E\53HUUHQVLQ*R\D&RUULHQWHV$UJHQWLQD name Marmosa marmota IRUWKHVDPHVSHFLPHQ
187 Gayana 73(2), 2009
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
FIGURA 3$ &RQVHQVRHVWULFWRGHiUEROHVPiVSDUVLPRQLRVRV ORQJLWXG SDVRV ,& ,5 REWHQLGRV HQHODQiOLVLVGHPi[LPDSDUVLPRQLDGHVHFXHQFLDVGHOJHQGHOFLWRFURPRE/RVQ~PHURVFRUUHVSRQGHQDYDORUHVGH DSR\RGH-DFNNQLIHGHORVQRGRVDVXGHUHFKD6yORVHPXHVWUDQYDORUHVGH-DFNNQLIHPD\RUHVD/RVQ~PHURVGH DFFHVRD*HQEDQNGHODVVHFXHQFLDVDQDOL]DGDVVHLQFOX\HQHQODHWLTXHWDVGHODVWHUPLQDOHV% &RQVHQVRSRUODUHJODGH ODPD\RUtDUHVXOWDGRGHODQiOLVLV%D\HVLDQRGHVHFXHQFLDVGHOJHQGHOFLWRFURPRE/RVQ~PHURVFRUUHVSRQGHQDYDORUHV GHSUREDELOLGDGDSRVWHULRULGHORVQRGRVDVXGHUHFKD/RVQ~PHURVGHDFFHVRD*HQEDQNGHODVVHFXHQFLDVDQDOL]DGDV VHLQFOX\HQHQODVHWLTXHWDVGHODVWHUPLQDOHV
Subsequently, after studying additional new use citellus because it is the more familiar name specimens from Corrientes, Thomas (1912) (marmotaKDVQRWEHHQXVHGLQWKHWD[RQRPLF described Marmosa citella, including in this literature for decades), and because it is not VSHFLHVWKHVSHFLPHQSUHYLRXVO\LGHQWL¿HGE\KLP associated with any nomenclatural ambiguity as Micoureus griseus and Marmosa marmota (marmota was used by Tate for the species The name Marmosa marmota (Oken, 1816), as currently known as T. macrurus 7KHUHSODFHPHQW used by Tate (1933) is not available (ICZN 1956), of a traditionally used name by an obscure epithet but the epithet marmota is alleged to have been formerly associated with a different species would made available inadvertantly by Thomas (1896: needlessly confuse current usage, so we follow IRRWQRWH &UHLJKWRQ *DUGQHU WKHUHFRPPHQGDWLRQVRIWKH,&=1> $UWV $OWKRXJK&UHLJKWRQ *DUGQHU DUJXHG DQG @LQPDLQWDLQLQJcitellus as that citella Thomas, 1912 is as a junior synonym the appropriate name for the Mesopotamian form of Marmosa marmota Thomas 1896, we prefer to of T. pusillus (sensu lato
188 7D[RQRP\RIThylamys: TETA,P.ET AL.
TABLE IV. Morphological comparison of Thylamys citellus, Tpulchellus, and Tpusillus
TABLA IV. Comparación morfológica de Thylamys citellus, Tpulchellus y Tpusillus
Thylamys citellus Thylamys pulchellus Thylamys pusillus
Tail bicolor, with white distal tip Tail bicolor, including the tip Tail bicolor, including the tip
Dorsal color pattern tricolor, with a Dorsal color pattern tricolor, with a Dorsal color pattern tricolor, with a general cinnamon tint general grayish brown tint general dark-brown tint
Supraorbital beads usually well Supraorbital beads slightly to well Supraorbital beads slightly to well developed developed developed
Postorbital constriction well Postorbital constriction slightly to Postorbital constriction slightly to marked well marked well marked
3RVWRUELWDOSURFHVVHVZHOOGH¿QHG Postorbital processes usually Postorbital processes usually absent in adult individuals DEVHQWRUVOLJKWO\GH¿QHG RUVOLJKWO\GH¿QHG Incisive foramina large, with Incisive foramina anteriorly Incisive foramina anteriorly subparallel medial and lateral H[SDQGHGDQGQDUURZLQLWV H[SDQGHGDQGQDUURZLQLWVSRVWHULRU margins posterior portion portion 3 3 3 3 3 3 Stylar cusp C in M3 usually well Stylar cusp C in M3 slightly to Stylar cusp C in M3 usually absent or marked well marked indistinct
FIGURE 4. Dorsal, ventral, and lateral views of skull and labial view of the mandible of the holotypes of: a-c) Marmosa citella7KRPDV %0 DQGGI Marmosa janetta pulchella &DEUHUD 0/3; 6FDOH EDU PP
FIGURA 4. Vista dorsal, ventral y lateral del cráneo y vista labial de la mandíbula de los holotipos de: a-c) Marmosa citella7KRPDV DE\F %0 \GI Marmosa janetta pulchella &DEUHUD 0/3; (VFDOD PP
189 Gayana 73(2), 2009
EMENDED DIAGNOSIS. A member of the didelphid ¿QHGRUVDOSHODJHVKRZLQJDFRQVSLFXRXVWULFRORU genus Thylamys intermediate in size between T. SDWWHUQZLWKDJHQHUDOFLQQDPRQWLQW )LJ 0LG pusillus and T. macrurus, characterized by a unique dorsal region from the ears to the hind-limbs is Raw combination of morphological traits including a 8PEHUZKLOHWKHVLGHVDUH:RRG%URZQ7KHGRUVDO distinct tricolored pattern with a general cinnamon SHODJHLVORQJDQGGHQVH JXDUGKDLUV PP DUH WLQW WDLOORQJHUWKDQKHDGDQGERG\ELFRORUHG EURZQLVKGDUNHUDWWKHLUEDVHV FRYHUKDLUVDUHVKRUW SUR[LPDOO\ GDUNDERYHZKLWLVKEHORZ EXW (8-9 mm) and four-banded, with a grayish base (2/3), ZLWKWKHWHUPLQDOWRHQWLUHO\ZKLWH VNXOO followed by a narrow dark brown band, and creamy robust with zygomatic arches proportionally wide EURZQGLVWDOSRUWLRQVDQGGDUNWLSV7KHYHQWHUIURP UHODWLYHWRFUDQLDOOHQJWK VXSUDRUELWDOULGJHVZHOO the cheeks to the anus, is covered by entire creamy GHYHORSHG SRVWRUELWDOFRQVWULFWLRQZHOOPDUNHG ZKLWHKDLUV PP 7KHJXODUUHJLRQLVZKLWHRU incisive foramina large, with subparallel medial KDVD\HOORZLVKWLQWLQVRPHLQGLYLGXDOV7KHKHDG DQGODWHUDOPDUJLQV SUHPD[LOODU\PD[LOODU\ is paler, with the snout characterized by a distinct VXWXUHMXVWDERYHRUSRVWHULRUWRWKH,DOYHROXV GDUNOLQH7KHH\HULQJLVPRGHUDWHO\GHYHORSHG SRVWHURODWHUDOSDODWDOIRUDPLQDORQJ H[WHQGLQJ DQGGDUNEURZQH[WHQGLQJVOLJKWO\WRWKHQRVH7KH to, or surpassing, the protocone of M4 on each UKLQDULXPKDVWZRYHQWURODWHUDOJURRYHVÀDQNLQJ VLGH DQGWKLUGXSSHUPRODUZLWKDPDUNHGVW\ODU WKHPHGLDQVXOFXVRQHDFKVLGH7KHYLEULVVDHDUH FXVS& )LJV PRVWO\GDUNEURZQH[FHSWIRUWKHLQWHUUDPDOKDLUV DESCRIPTION Thylamys citellus is a large, long- (which are whitish) and some large mystacial hairs tailed mouse opossum, with a brownish, soft, and WKDWKDYHZKLWLVKGLVWDOSRUWLRQV7KHHDUVDUH
FIGURE 5'RUVDODQGYHQWUDOYLHZVRIVWXG\VNLQVRIDG Thylamys citellus (MACN 16262: Argentina, Entre Ríos, 3URQXQFLDPLHQWR EH Tpulchellus 0$&1 $UJHQWLQD6DQWLDJRGHO(VWHUR6DQ$QWRQLR DQGFI Tpusillus 0$&1 $UJHQWLQD)RUPRVD/DJXQD%ODQFD
FIGURA 59LVWDVGRUVDO\YHQWUDOGHSLHOHVGHHVWXGLRGHDG Thylamys citellus (MACN 16262: Argentina, Entre Ríos, 3URQXQFLDPLHQWR EH Tpulchellus (MACN 17279: Argentina, Santiago del Estero, San Antonio) y c, f) Tpusillus 0$&1 $UJHQWLQD)RUPRVD/DJXQD%ODQFD
190 7D[RQRP\RIThylamys: TETA,P.ET AL. large and rounded, partially covered by very short WDLOLVHQWLUHO\ZKLWH&DXGDOVFDOHVDUHDUUDQJHGLQ grayish-brown hairs, and apparently naked without DQQXODUVHULHV7KHYHQWUDOVXUIDFHRIWKHWDLOWLS PDJQL¿FDWLRQ7KHVNLQRIWKHHDUVLVGDUNEURZQ PP LVPRGL¿HGIRUSUHKHQVLRQZLWKDQDNHGPHGLDQ DWWKHWLSDQG\HOORZLVKDWWKHEDVH%HKLQGHDFK JURYHDQGDÀHVK\DSLFDOSDGEHDULQJGHUPDWRJO\SKV HDUWKHUHLVDFRQVSLFXRXVWXIWRIZKLWLVKKDLUV Females do not have a pouch, and 13 mammae are The fore and hind limbs are whitish, without color arranged in a nearly circular pattern in the abdomen differentiation between the dorsum and the palmar 7KHVNXOOLVUREXVWZLWKZLGH]\JRPDWLF DQGSODQWDUVXUIDFHV7KHIHHWDUHODUJHDQGDOOWKH arches relative to cranial length (the average ratio WRHVKDYHODUJHZKLWHXQJXDOWXIWV7KHWRHVDUH of zygomatic breadth to condylobasal length × 100 long, with short claws that do not surpass the apical LVDERXW )LJ 7KHLQWHURUELWDOUHJLRQ pads on the forefeet, but are slightly longer on the is moderately narrowed, and mature adults often KLQGIHHW%RWKPDQXVDQGSHVKDYHVL[ODUJHQHDUO\ have pointed postorbital processes from which RYRLGVHSDUDWHSDGV3DOPDUDQGSODQWDUVXUIDFHVDUH VKDUSULGJHVFRQYHUJHSRVWHULRUO\RQWRWKHSDULHWDOV FRYHUHGZLWKODUJHJUDQXOHVDQGGHUPDWRJO\SKV7KH 7KHSRVWRUELWDOFRQVWULFWLRQLVZHOOPDUNHG7KH tail is long, slightly incrassated in some individuals braincase is inflated, sometimes with weakly (6-7 mm in diameter), apparently naked without GH¿QHGWHPSRUDOULGJHV7KHODPEGRLGDOFUHVWVDUH PDJQL¿FDWLRQDQGELFRORUHG7KHEDVHRIWKHWDLO FD VOLJKWO\RUPRGHUDWHO\GHYHORSHG,QODWHUDOYLHZ PP LVVOLJKWO\FRYHUHGE\WKHERG\SHODJH7KH WKHSUHPD[LOODU\PD[LOODU\VXWXUHLVSRVWHULRUWRWKH tail dorsum is brownish, sharply demarcated from ,DOYHROXV7ZRODFULPDOIRUDPLQDDUHFRQWDLQHG WKHZKLWLVKYHQWHU7KHGLVWDOWLS WR RIWKH within the anterior orbital margin, being small and
FIGURE 6. Dorsal and ventral views of the skulls of: a, d) Thylamys pulchellus (MACN 17279: Argentina, Santiago del (VWHUR6DQ$QWRQLR EH Tpusillus )01+ 3DUDJXD\%RTXHUyQ6FKPLGW5DQFKNPHDVW&RUUDOHV DQG c, f) Tcitellus 0$&1 $UJHQWLQD(QWUH5tRV3DUTXH1DFLRQDO(O3DOPDU 6FDOHEDU PP
FIGURA 6. Vista dorsal y ventral de los cráneos de: a, d) Thylamys pulchellus (MACN 17279: Argentina, Santiago del (VWHUR6DQ$QWRQLR EH Tpusillus (FMNH 164086: Paraguay, Boquerón, Schmidt Ranch, 10 km east Corrales) y c, f) Tcitellus 0$&1 $UJHQWLQD(QWUH5tRV3DUTXH1DFLRQDO(O3DOPDU (VFDOD PP
191 Gayana 73(2), 2009
VOLJKWO\YLVLEOHLQODWHUDOYLHZ7KHQDVDOVDUHORQJ with subparallel lateral margins, and have a slight H[SDQVLRQDWWKHPD[LOORIURQWDOVXWXUHQDUURZLQJ VOLJKWO\EHKLQGLW7KHLQIUDRUELWDOIRUDPHQLVODUJH DQGLVORFDWHGMXVWDERYH37KH]\JRPDWLFDUFKHV DUHFRQVSLFXRXVO\H[SDQGHGDQGWKHH[WHUQDOVXUIDFH RIWKHMXJDOVH[KLELWDFRQVSLFXRXVFRQFDYLW\DORQJ WKHLUDQWHULRUSRUWLRQ7KHLQFLVLYHIRUDPLQDDUHODUJH with subparallel medial and lateral margins, reaching LQLWVSRVWHULRUH[WHQVLRQWKHPLGOLQHEHWZHHQ FDQLQHV7KHSDODWHLVKLJKO\IHQHVWUDWHGVKRZLQJ PD[LOORSDODWLQHSDODWLQHDQGPD[LOODU\RSHQLQJV 7KHPD[LOORSDODWLQHIHQHVWUDHDUHODUJHDQGZLGH H[WHQGLQJRQHDFKVLGHIURPWKHDQWHULRUSRUWLRQ RI3WRWKHDQWHULRUSRUWLRQRI07KHPD[LOODU\ fenestrae are small (sometimes unilaterally present) DQGDUHORFDWHGEHWZHHQ0DQGWKHPD[LOORSDODWLQH IHQHVWUDH7KHSRVWHURODWHUDOSDODWDOIRUDPLQDDUH large and wide, reaching or surpassing the lingual DSLFHVRIWKHSURWRFRQHVRI07KHDOLVSKHQRLG W\PSDQLFZLQJVDUHVPDOODQGDOPRVWVSKHULFDO7KH I2 to I5 are subequal in size, nearly rhomboidal, and LQFUHDVLQJLQEUHDGWKIURPIURQWWREDFN&DQLQHVDUH ODUJHDQGODFNDFFHVVRU\FXVSV3LVODUJHUWKDQ3 P1 is present and smaller than other premolars but QRWYHVWLJLDO7KHXSSHUPRODUVDUHGLODPERGRQW compressed antero-posteriorly (especially M4), and highly carnassialized, increasing in width from 0WR07KHHFWRÀH[XVLVDEVHQWRQ0YHU\ FIGURE 72FFOXVDOVXUIDFHRIWKHULJKWXSSHUPRODUVHULHV VKDOORZRQ0DQGGLVWLQFWRQ06W\ODUFXVS& of: a) Thylamys citellus (MACN 23459: Argentina, Entre is well developed on the upper molars, being similar 5tRV3DUTXH1DFLRQDO(O3DOPDU E Tpulchellus (MACN in size to stylar cusps B and D in M1, and smaller $UJHQWLQD6DQWLDJRGHO(VWHUR6DQ$QWRQLR F WKDQVW\ODUFXVSV%DQG'LQ0DQG0 )LJ Tpusillus (MACN 20779: Argentina, Formosa, Laguna %ODQFD DQGG Tpusillus (FMNH 164086: Paraguay, The horizontal and ascending rami of the mandible Boquerón, Schmidt Ranch, 10 km east Corrales) showing IRUPDQRSHQDQJOH7KHORZHULQFLVRUVKDYH the level of development of stylar cusp C (indicated by GLVWLQFWOLQJXDOFXVSV$VPDOOVRPHWLPHVLQGLVWLQFW WKHDUURZ $EEUHYLDWXUHV 33 XSSHUVHFRQGDQGWKLUG SRVWHULRUFXVSLVSUHVHQWRQF7KHVHFRQGDQGWKLUG SUHPRODU 0000 XSSHU¿UVWVHFRQGWKLUGDQG ORZHUSUHPRODUV SDQGS DUHVXEHTXDOLQVL]H IRXUWKPRODU P PD[LOODU\IHQHVWUD PS PD[LOORSDODWLQH IHQHVWUD SOSI SRVWHURODWHUDOSDODWDOIRUDPHQ 7KHWU\JRQLGLQFUHDVHVLQOHQJWKIURPPWRP WKH WDORQLGLVQHDUO\VTXDUHG7KHK\SRFRQLGRQPLV FIGURA 7.6XSHU¿FLHRFOXVDOGHODVHULHPRODUVXSHULRU QRWODELDOO\VDOLHQW WKHHQWRFRQLGLVODUJHDQGZHOO izquierda de: a) Thylamys citellus (MACN 23459: GHYHORSHGRQPP2EVHUYHGJHQHWLFGLVWDQFH $UJHQWLQD(QWUH5tRV3DUTXH1DFLRQDO(O3DOPDU E among studied haplotypes of the cytochrome b gene Tpulchellus (MACN 17279: Argentina, Santiago del (VWHUR6DQ$QWRQLR F Tpusillus (MACN 20779: ¿UVWES UHFRYHUHGIURPWKUHHVSHFLPHQVRIT. Argentina, Formosa, Laguna Blanca) y d) Tpusillus citellusLV (FMNH 164086: Paraguay, Boquerón, Schmidt Ranch, 10 km east Corrales) mostrando el nivel de desarrollo de COMPARISONS. The known geographic range of T. ODF~VSLGHHVWLODU& LQGLFDGDFRQODÀHFKD $EUHYLDWXUDV citellus is adjacent to the distributions of three 33 VHJXQGR\WHUFHUSUHPRODUVXSHULRU 000 0 SULPHURVHJXQGRWHUFHUR\FXDUWRPRODUVXSHULRU congeneric species, T. macrurus, T. pusillus, and T. P IHQHVWUDPD[LODU PS IHQHVWUDPD[LORSDODWLQD SOSI pulchellus )LJ Thylamys macrurus is larger than IRUDPHQSDODWDOSRVWHURODWHUDO T. citellus, especially in molar dimensions (length
192 7D[RQRP\RIThylamys: TETA,P.ET AL.
RIXSSHUPRODUURZ PPYVPP Thylamys pulchellus (Cabrera, 1934) Table II) and has much smaller posterolateral )LJV7DEOH,, ,9 SDODWDOIRUDPLQD VHH&DUPLJQRWWR 0RQIRUW )LJ Thylamys pusillus is slightly smaller HOLOTYPE. 0/3; DGXOWIHPDOH FROOHFWHG (intermediate in size between T. citellus and T. by Jorge Argañaraz and donated to MLP by Ángel pulchellus), lacks a pale tail-tip, has a darker &DEUHUD &DEUHUD )LJ dorsal coloration, and usually lacks well developed VXSUDRUELWDOULGJHVDQGVW\ODUFXVS&RQ0 )LJ TYPE LOCALITY. “Los Robles, Santiago del Estero” 7DEOH,9 Thylamys pulchellus is much smaller &DEUHUD )LJ than T. citellus (length of upper molar row usually PP 7DEOH,, KDVJUD\LVKEURZQGRUVDO TAXONOMIC REMARKS7KHW\SHPDWHULDORIpulchellus SHODJHDQGODFNVDSDOHWDLOWLS$GGLWLRQDOO\WKH is from Santiago del Estero Province in the western SUHPD[LOODU\PD[LOODU\VXWXUHLQERWKTpulchellus $UJHQWLQHDQSRUWLRQRIWKH'U\&KDFRHFRUHJLRQ and Tpusillus is anterior to the I5 alveolus, while Voss et al. LQSUHVV VHHDOVR9RVV -DQVD in TcitellusWKLVVXWXUHLVSRVWHULRUWR, )LJ suggested that an older available name for this 7DEOH,9 2EVHUYHGJHQHWLFGLVWDQFHLQWKH¿UVW species could be bruchi Thomas, 1921, with type bp of the cytochrome b gene between T. citellus ORFDOLW\LQ$OWR3HQFRVRQRUWKRI6DQ/XLV3URYLQFH and T. pulchellusLVDQGEHWZHHQT. citellus Although bruchi was included in the synonymy of and T. pusillusLV )LJ Thylamys pallidior by Gardner (2005) and Creighton *DUGQHU DFXUVRU\H[DPLQDWLRQRIWKHW\SH MEASUREMENTS. ([WHUQDO DQG FUDQLRGHQWDO specimen of bruchi, housed at the British Museum, measurements of T. citellus are provided in Table UHYHDOVWKHSUHVHQFHRIGLVWLQFWPD[LOODU\YDFXLWLHV ,, and well developed stylar cusp C on M1 and M2 that better match with the diagnostic traits of T. HABITAT AND DISTRIBUTION. T. citellus is found in pulchellus (as recognized herein) than with those the Southern Cone Mesopotamian Savanna and of Tpallidior+RZHYHUbruchiODFNVZHOOGH¿QHG the Mesopotamian sector of the Humid Pampas supraorbital borders, and has large auditory bullae, eco-regions (sensu Olson y Dinerstein 2002) in the resembling those of T. pallidior, so we provisionally Argentine provinces of Entre Ríos and Corrientes follow the opinion of Tate (1933), who allied SURYLQFHV )LJ 7KHODQGVFDSHLQWKLVDUHDLVD bruchi with pallidior on the basis of its coloration, PL[WXUHRIJUDVV\VDYDQQDVRSHQJUDVVODQGVDQG ERG\SURSRUWLRQVDQGVRPHFUDQLDOIHDWXUHV HJ vast wetlands with patches of subtropical gallery GHYHORSPHQWRIWKHDXGLWRU\EXOODH 3DUWRIWKLV IRUHVW[HURSK\WLFZRRGODQGVDQGSDOPVDYDQQDV confusing situation is due to the fact that the type The region is highly threatened due to destruction specimen of bruchi is a young individual without the and degradation of the natural habitat by housing, FRPSOHWHGHQWLWLRQHUXSWHG,WLVDOVRUHOHYDQWWKDW cattle ranching and agriculture (Olson et al. RWKHUH[DPLQHGVSHFLPHQVIURP6DQ/XLV3URYLQFH The known distributional range of T. citellus is including some from near the type locality of bruchi, limited by the Paraná River to the north and west, FOHDUO\¿WWKHGLDJQRVLVRIT pallidior$GGLWLRQDO DQGE\WKH8UXJXD\5LYHUDQGWKHÀRRGHGJUDVVODQGV genetic and morphological evidence from topotypic RIWKH,EHUiZHWODQGVWRWKHHDVW1RRWKHUThylamys material of bruchiDVZHOODWKRURXJKH[DPLQDWLRQ species is sympatric with T. citellus*RQ]DOH]et of the type specimen is highly necessary to assess al. (2000) reported a specimen of Thylamys from WKHVWDWXVRIWKLVQRPLQDOWD[RQ Uruguay (east of the Uruguay River), but they GLGQRWLGHQWLI\LWWRWKHVSHFLHVOHYHO:HGLGQRW EMENDED DIAGNOSIS. A small member of the directly study this Uruguayan specimen, but an didelphid genus Thylamys, characterized by a unique LQVSHFWLRQRIWKH¿JXUHSURYLGHGE\*RQ]DOH]et combination of morphological traits including a al. (2000) suggests it may not belong in Thylamys moderately marked tricolored pattern with a general because its nasals are not parallel-sided and have JUD\LVKEURZQDSSHDUDQFH WDLOORQJHUWKDQKHDGDQG DFRQVSLFXRXVH[SDQVLRQDWWKHLUSRVWHULRUSRUWLRQ body and sharply bicolored (dark above, whitish (Gonzalez et al. )LJ $GLUHFWLQVSHFWLRQ EHORZ VNXOOVPDOOEXWVWURQJO\EXLOW VXSUDRUELWDO RIWKHVSHFLPHQZRXOGFODULI\WKLVLVVXH ULGJHVVOLJKWO\WRZHOOGHYHORSHG SRVWRUELWDO
193 Gayana 73(2), 2009
FRQVWULFWLRQZHOOPDUNHG LQFLVLYHIRUDPLQDDQWHULRUO\ indistinct in some individuals but well marked in H[SDQGHGDQGQDUURZLQWKHLUSRVWHULRUSRUWLRQ RWKHUV7KHEUDLQFDVHLVLQÀDWHGVRPHWLPHVZLWK SUHPD[LOODU\PD[LOODU\VXWXUHDQWHULRURUMXVWDERYH ZHDNO\GH¿QHGWHPSRUDOULGJHV7KHODPEGRLGDO ,DOYHROXV ORQJSRVWHURODWHUDOSDODWDOIRUDPLQD FUHVWVDUHLQFRQVSLFXRXV,QODWHUDOYLHZWKH H[WHQGLQJWRRUVXUSDVVLQJWKHSURWRFRQHRI0 SUHPD[LOODU\PD[LOODU\VXWXUHLVDQWHULRUWRWKH RQHDFKVLGH DQGWKLUGXSSHUPRODUZLWKDPDUNHG ,DOYHROXV7KHODFULPDOIRUDPLQDDUHFRQWDLQHG VW\ODUFXVS& within the anterior orbital margin, being usually ODUJHDQGZHOOYLVLEOHLQODWHUDOYLHZ7KHQDVDOV DESCRIPTION. Thylamys pulchellus is a small, long- are long, with subparallel lateral margins, and have WDLOHGPRXVHRSRVVXPZLWKVRIW¿QHJUD\LVK DVOLJKWH[SDQVLRQDWWKHPD[LOORIURQWDOVXWXUH brown dorsal pelage showing a moderately marked QDUURZLQJVOLJKWO\EHKLQGLW7KHLQIUDRUELWDO WULFRORUSDWWHUQ )LJ 7KHPLGGRUVDOIXUIURP IRUDPHQLVODUJHDQGLVORFDWHGMXVWDERYH37KH WKHHDUVWRWKHKLQGOLPEVLV:DUP6HSLDWR6HSLD H[WHUQDOVXUIDFHRIWKHMXJDOVH[KLELWVDFRQVSLFXRXV ZKLOHWKHÀDQNVDUH+DLU%URZQ&RYHUKDLUV FRQFDYLW\DORQJWKHLUDQWHULRUSRUWLRQ7KHLQFLVLYH mm) are four banded, with gray bases (4/5), middle IRUDPLQDDUHODUJHDQWHULRUO\H[SDQGHGDQGQDUURZ portions dark brown to creamy brown, and dark in their posterior portion, reaching the anterior line WLSV9HQWUDOKDLUVIURPWKHFKHHNVWRWKHDQXV EHWZHHQFDQLQHV7KHSDODWHLVKLJKO\IHQHVWUDWHG DUHHQWLUHO\FUHDP\ZKLWH PP 7KHKHDGLV VKRZLQJPD[LOORSDODWLQHSDODWLQHDQGPD[LOODU\ paler, with the snout characterized by a distinct RSHQLQJV7KHPD[LOORSDODWLQHIHQHVWUDHDUHODUJH GDUNOLQH7KHH\HULQJLVPRGHUDWHO\GHYHORSHG DQGZLGHH[WHQGLQJRQHDFKVLGHIURPWKHFRPPL DQGGDUNEURZQH[WHQGLQJVOLJKWO\WRWKHQRVH7KH sure between P2 and P3 to the middle portion of UKLQDULXPKDVWZRYHQWURODWHUDOJURRYHVÀDQNLQJ 07KHPD[LOODU\IHQHVWUDHDUHODUJHH[WHQGLQJ WKHPHGLDQVXOFXVRQHDFKVLGH7KHYLEULVVDHDUH IURPWKH3WRWKH00FRPPLVVXUH7KH PRVWO\GDUNEURZQH[FHSWIRUWKHLQWHUUDPDOKDLUV posterolateral palatal foramina are large and wide, (which are whitish), and some large mystacial surpassing the lingual apices of the protocone of KDLUVWKDWKDYHZKLWLVKGLVWDOSRUWLRQV7KHHDUV 0RQHDFKVLGH7KHDOLVSKHQRLGW\PSDQLFZLQJV are large and rounded, entirely covered by very DUHVPDOODQGDOPRVWVSKHULFDO7KHXSSHULQFLVRUV short grayish-brown hairs and apparently naked (I2-I5) are subequal in size, nearly rhomboidal, ZLWKRXWPDJQL¿FDWLRQ7KHIRUHDQGKLQGOLPEVDUH DQGLQFUHDVLQJLQEUHDGWKIURPIURQWWREDFN7KH whitish, without color differentiation between the FDQLQHVDUHODUJHDQGODFNDFFHVVRU\FXVSV7KH GRUVXPDQGWKHSDOPDUDQGSODQWDUVXUIDFHV7KH WKLUGXSSHUSUHPRODU 3 LVVOLJKWO\ODUJHUWKDQ3 feet are small, and all the toes have large, white, P1 is present and smaller than other premolars but XQJXDOWXIWV7KHWRHVDUHORQJZLWKVKRUWFODZV QRWYHVWLJLDO7KHXSSHUPRODUVDUHGLODPERGRQW that do not surpass the apical pads in forefeet, but compressed antero-posteriorly (especially the M4), DUHVOLJKWO\ORQJHULQWKHKLQGIHHW%RWKPDQXV and highly carnassialized, increasing in width from DQGSHVKDYHVL[ODUJHRYRLGVHSDUDWHSDGV 0WR07KHHFWRÀH[XVLVDEVHQWRQ0YHU\ The tail is long, slightly incrassated (6-7 mm in VKDOORZRQ0DQGGLVWLQFWRQ06W\ODUFXVS& diameter) in some individuals, apparently naked is well developed on M1-M3, although it is smaller ZLWKRXWPDJQL¿FDWLRQDQGDOPRVWELFRORUHG7KH WKDQVW\ODUFXVSV%DQG' )LJ 7KHKRUL]RQWDO tail dorsum is brownish, sharply demarcated from and ascending rami of the mandible form an open WKHZKLWLVKYHQWHU&DXGDOVFDOHVDUHDUUDQJHGLQ DQJOH7KHORZHULQFLVRUVKDYHLQGLVWLQFWOLQJXDO DQQXODUVHULHV7KHYHQWUDOVXUIDFHRIWKHWDLOWLS FXVSV7KHORZHUFDQLQHKDVDVPDOOXVXDOO\ZHOO PP LVPRGL¿HGIRUSUHKHQVLRQZLWKD PDUNHGSRVWHULRUFXVS7KHVHFRQGDQGWKLUG QDNHGPHGLDQJURYHDQGDÀHVK\DSLFDOSDGEHDULQJ ORZHUSUHPRODUVDUHVXEHTXDOLQVL]H7KHWULJRQLG GHUPDWRJO\SKV)HPDOHVGRQRWKDYHDSRXFK7KH LQFUHDVHVLQOHQJWKIURPPWRP WKHWDORQLG skull is small but strongly built, with wide zygomatic LVQHDUO\VTXDUHG7KHK\SRFRQLGRQPLVQRW arches relative to cranial length (the average ratio ODELDOO\VDOLHQW WKHHQWRFRQLGLVODUJHDQGZHOO of zygomatic breadth to condylobasal length × 100 GHYHORSHGRQPP2EVHUYHGJHQHWLFGLVWDQFH LVDERXW )LJ 7KHLQWHURUELWDOUHJLRQ DPRQJVWXGLHGKDSORW\SHVFRUUHVSRQGLQJWRWKH¿UVW is narrow, and the supraorbital margins are slightly 801 bp of the cytochrome b gene recovered from WRGLVWLQFWO\EHDGHG7KHSRVWRUELWDOFRQVWULFWLRQLV specimens of T. pulchellusLV 7KHNDU\RW\SH
194 7D[RQRP\RIThylamys: TETA,P.ET AL. is 2n )1 WKH;FKURPRVRPHLVDVPDOO DISCUSSION submetacentric and the Y chromosome is missing in somatic cells (Braun et alE The number of recognized species within Thylamys KDVYDULHGVLJQL¿FDQWO\GXULQJWKHODVWWZRGHFDGHV COMPARISONS. The known geographic range of )RULQVWDQFH*DUGQHU UHFRJQL]HGRQO\¿YH Thylamys pulchellus is adjacent to the distributions species, while in the last treatise of South American of three congeneric species, Tcitellus, T. pallidior, PDUVXSLDOV&UHLJKWRQ *DUGQHU UHFRJQL]H and T. pusillus )LJ T. citellus is clearly larger EXWVHH9RVV -DQVD $VFXUUHQWO\UHFRJQL]HG than T. pulchellus, has a more cinnamon general T. pusillusLVDFRPSOH[RIDWOHDVWWKUHHGLDJQRVDEOH DSSHDUDQFHKDVDZKLWHWDLOWLS )LJ7DEOH WD[DWZRRIZKLFK SUHYLRXVO\WUHDWHGDVV\QRQ\PV ,9 Thylamys pusillus is slightly larger than T. of pusillus) are elevated to species level in this pulchellus, has darker fur, frequently lacks stylar UHSRUW7KLVWD[RQRPLFUHVROXWLRQDWOHDVWLQSDUW FXVS&RQ0 )LJ DQGXVXDOO\KDVPXFK was previously suggested by Voss et al. (in press), VPDOOHUPD[LOODU\YDFXLWLHV2EVHUYHGJHQHWLF who reported some metric and dental differences GLVWDQFHLQWKH¿UVWESRIWKHF\WRFKURPHE between typical Paraguayan samples of T. pusillus gene between T. pulchellus and T. pusillusLV (sensu stricto) and the Argentinean populations herein )LJ Thylamys pallidior has much longer fur referred to Tpulchellus,QDEURDGHUFRQWH[WRXU (usually >10 mm) than T. pulchellus, has longer ¿QGLQJVDUHLQOLQHZLWKRWKHUUHFHQWDQGSDUWLDOO\ PDQXDOFODZV DOZD\VH[WHQGLQJEH\RQGWKHÀHVK\ XQH[SHFWHGGLVFRYHULHVZLWKLQWKH'LGHOSKLGDHVXFK apical pad of each digit), larger bullae, consistently as the recent description of several cryptic species, ODFNVPD[LOODU\IHQHVWUDHDQGODFNVDQGVW\ODUFXVS and even genera (Flores et al.9RVV -DQVD &RQ00 9RVVet al.DE
MEASUREMENTS. ([WHUQDO DQG FUDQLRGHQWDO *LYHQRXUWD[RQRPLFUHVXOWVWKHJHQXVThylamys is measurements of T. pulchellus are provided in now composed of 11 species, of which 6 occur in 7DEOH,, $UJHQWLQDDQGDUH$UJHQWLQHDQHQGHPLFV)XWXUH studies should refine species boundaries within HABITAT AND DISTRIBUTION. T. pulchellus is endemic Thylamys)RULQVWDQFHLWVHHPVZRUWKLQYHVWLJDWLQJWKH to the Dry Chaco eco-region (sensu Olson y degree of geographic differentiation within T pallidior, Dinerstein 2002) of the provinces of Catamarca, another widely distributed species with populations Chaco, San Juan, Salta, and Santiago del Estero ranging from northern Chile to Argentinean Patagonia )LJ DQGSHUKDSVDOVRRFFXUVLQ)RUPRVDDQG &UHLJKWRQ *DUGQHU DQGDVSHFLHVWKDWDOVR Tucumán (see Flores et al 7KHODQGVFDSH VKRZVSK\ORJHRJUDSKLFVWUXFWXUH )LJ $SSHQGL[ LQWKLVELRPHLVDPL[WXUHRIJUDVVODQGVDQG see also Braun et al.D [HURSK\WLFZRRGODQGVQRZW\SLFDOO\GRPLQDWHGE\ secondary growth forests of Schinopsis lorentzii, Thylamys citellus, Tpusillus, and T. pulchellus form a Aspidosperma quebracho-blanco, and Prosopis VWURQJO\VXSSRUWHGFODGH )LJ ZKLFKVXJJHVWVWKDW VS0RUSKRPHWULFDQGPRUSKRORJLFDOHYLGHQFHVX SDUWRIWKHGLYHUVL¿FDWLRQRIThylamys has occurred ggests that both pulchellus and pusillus are present LQVLGHWKH/D3ODWD5LYHUEDVLQ3ODXVLEO\FHUWDLQ LQ$UJHQWLQD+RZHYHUWKHLUGLVWULEXWLRQDOERXQ natural geographical barriers such as major rivers might GDULHVDUHXQFOHDU VHHDERYH ,WLVQRWFXUUHQWO\ have played a strong role delimiting local species of known whether or not these species occur in Thylamys: the sister species T. citellus and T. pulchellus, sympatry along the transitional zone between are geographically separated by the large Paraná River, the Dry and Humid Chaco (the latter inhabited IRUH[DPSOHZKHUHDV T. pulchellus and T. pusillus by T. pusillus $WWKHVDPHWLPHWKHJHRJUDSKLF VHHPVWREHVHSDUDWHGE\WKH%HUPHMR5LYHU6LPLODUO\ distribution of T. pulchellus to the west and south T. pusillus does not cross the Paraguay River and is (Córdoba and Catamarca Provinces) remains WKXVUHVWULFWHGWRWKH'U\&KDFR$GGLWLRQDOVWXGLHV XQFOHDU,QWKRVHDUHDVLWFRXOGRYHUODSZLWKT. in line to those of Patton et al. (1994, 2000), together pallidior (Thomas, 1902), from which it is clearly ZLWKIXUWKHU¿HOGFROOHFWLRQVDUHQHHGHGWRFODULI\LI differentiable both morphologically and genetically WKHVHULYHUVSURPSWHGWKHGLYHUVL¿FDWLRQRIThylamys )LJ RUMXVWRSHUDWHDVGLVSHUVDOEDUULHUV
195 Gayana 73(2), 2009
$&.12:/('*0(176 *UD\,Q 0DPPDOVRI6RXWK$PHULFD 9ROXPH,0DUVXSLDOV;HQDUWKUDQV6KUHZVDQG Robert Baker (TTU), Bruce Patterson (FMNH), %DWV (G*DUGQHU$/,Q*DUGQHU$/ SS 8QLYHUVLW\RI&KLFDJR3UHVV&KLFDJR .XUW6FKZHUN 8&21 5XEpQ%DUTXH] &0/ DQG/RQGRQ Ulyses Pardiñas (CNP), and Diego Verzi (MLP) D’ELÍA, G. & PARDIÑAS, U.F.J. 2004. Systematics of kindly permitted us the study of Thylamys specimens Argentinean, Paraguayan, and Uruguayan XQGHUWKHLUFDUH5REHUW9RVVDQG6HUJLR6RODUL swamp rats of the genus Scapteromys (Rodentia, made valuable comments that greatly improved the &ULFHWLGDH 6LJPRGRQWLQDH -RXUQDO RI 0DPPDORJ\ TXDOLW\RIWKLVPDQXVFULSW:HDUHDOVRJUDWHIXOWR FARRIS, J.S. 7KHORJLFDOEDVLVRISK\ORJHQHWLF Isabel Gómez Villafañe and Ulyses Pardiñas, who DQDO\VLV,Q $GYDQFHVLQ&ODGLVWLFV 3URFHHGLQJV generously shared some specimens collected by RIWKH6HFRQG0HHWLQJRIWKH:LOOL+HQQLJ6RFLHW\ WKHLUUHVHDUFKWHDPVLQ(QWUH5tRVSURYLQFH6SH- (GV3ODQLFN1 )XQN9 SS&ROXPELD 8QLYHUVLW\3UHVV1HZ
196 7D[RQRP\RIThylamys: TETA,P.ET AL.
REIG, O. A., KIRSCH, J.A.W. & MARSHALL, L.G. HUELSENBECK, J. P., RONQUIST, F., NIELSEN, R. & BOLLBACK, Systematic relationships of the living and J.P.%D\HVLDQLQIHUHQFHRISK\ORJHQ\DQG 1HRFHQR]RLF$PHULFDQ µµRSRVVXPOLNH¶¶ LWVLPSDFWRQHYROXWLRQDU\ELRORJ\6FLHQFH marsupials (suborder Didelphimorphia), with comments on the classification of these and ,&=1 ,QWHUQDWLRQDOFRGHRI]RRORJLFDOQRPHQFODWXUH RIWKH&UHWDFHRXVDQG3DOHRJHQH1HZ:RUOG WKHG/RQGRQ ,QWHUQDWLRQDO7UXVWIRU=RRORJLFDO DQG(XURSHDQPHWDWKHULDQV,Q 3RVVXPVDQG 1RPHQFODWXUH RSRVVXPV VWXGLHVLQHYROXWLRQ (G$UFKHU JANSA, S.A. & VOSS, R.S. 20003K\ORJHQHWLFVWXGLHV 0 YROSS 6\GQH\ 6XUUH\%HDWW\DQG RQGLGHOSKLGPDUVXSLDOV,,QWURGXFWLRQDQG 6RQV preliminary results from nuclear IRBP gene RIDGWAY,R. &RORUVWDQGDUGVDQGFRORUQRPHQFODWXUH VHTXHQFHV-RXUQDORI0DPPDOLDQ(YROXWLRQ :DVKLQJWRQ'& $XWKRUSS RONQUIST, F. & HUELSENBECK, J.P.05%$<(6 KIRSCH, J.A.W. & PALMA, R.E. 1 '1$'1$ %D\HVLDQSK\ORJHQHWLFLQIHUHQFHXQGHUPL[HG K\EULGL]DWLRQVWXGLHVRIFDUQLYRURXVPDUVXSLDOV PRGHOV%LRLQIRUPDWLFV 9$IXUWKHUHVWLPDWHRIUHODWLRQVKLSVDPRQJ SOLARI,S.'LYHUVLW\DQGGLVWULEXWLRQRIThylamys RSRVVXPV 0DUVXSLDOLD 'LGHOSKLGDH 0DPPDOLD (Didelphidae) in South America, with emphasis RQVSHFLHVIURPWKHZHVWHUQVLGHRIWKH$QGHV MEYNARD, A.P., PALMA, R.E. & RIVERA-MILLA,( In: Predators with pouches – the biology of )LORJHRJUDItDGHODVOODFDVFKLOHQDVGHOJpQHUR FDUQLYRURXVPDUVXSLDOV -RQHV0'LFNPDQ Thylamys (Marsupialia, Didelphidae) en base & $UFKHU0HGV SS&6,52 DVHFXHQFLDVGHOJHQPLWRFRQGULDOFLWRFURPRE 3XEOLVKLQJ0HOERXUQH Revista Chilena de Historia Natural 79(2):299- STATSOFT,INC6WDWLVWLFDIRU:LQGRZV FRPSXWHU SURJUDP PDQXDO 6WDW6RIW ,QF 7XOVD OLSON, D.M., DINERSTEIN, E., WIKRAMANAYAKE, E.D., 2NODKRPD BURGESS,N.D., POWELL, G.V.N., UNDERWOOD, STEINER, C., TILAK, M., DOUZERY, E.J.P. & CATZEFLIS, E.C., D’AMICO, J.A., STRAND, H.E., MORRISON, F.M.1HZ'1$GDWDIURPDWUDQVWK\UHWLQ J.C., LOUCKS, C.J., ALLNUTT, T.F., LAMOREUX, J.F., nuclear intron suggest an Oligocene to Miocene RICKETTS, T.H., ITOUA, I., WETTENGEL, W.W, KURA, diversification of living South American Y. & KASSEM,H.7HUUHVWULDOHFRUHJLRQVRI RSRVVXPV 0DUVXSLDOLD 'LGHOSKLGDH 0ROHFXODU WKHZRUOG DQHZPDSRIOLIHRQ(DUWK%LR6FLHQFH 3K\ORJHQHWLFVDQG(YROXWLRQ SWOFFORD, D.L.3$83 3K\ORJHQHWLF$QDO\VLV OLSON, D.M. & DINERSTIEN, E.7KH*OREDO XVLQJ3DUVLPRQ\ DQGRWKHUPHWKRGV 6LQDXHU 3ULRULW\HFRUHJLRQVIRUJOREDOFRQVHUYDWLRQ $VVRFLDWHV6XQGHUODQG0DVVDFKXVHWWV Annals of the Missouri Botanical Garden TAMURA, K., DUDLEY, J., NEI, M. & KUMAR,S.0(*$ 4: Molecular Evolutionary Genetics Analysis PALMA, R.E. & YATES, T.L. 1998. Phylogeny of southern 0(*$ VRIWZDUHYHUVLRQ0ROHFXODU%LRORJ\ South American mouse opossums (Thylamys, DQG(YROXWLRQ Didelphidae) based on allozyme and chromosomal TATE, G.H.H. $V\VWHPDWLFUHYLVLRQRIWKHPDUVXSLDO GDWD=HLWVFKULIWIU6DXJHWLHUNXQGH genus Marmosa, with a discussion of the adaptive PALMA, R.E., RIVERA-MILLA, E., YATES, T.L., MARQUET, UDGLDWLRQRIWKHPXULQHRSRVVXPV%XOOHWLQRIWKH P.A. & MEYNARD, A.P. 3K\ORJHQHWLFDQG $PHULFDQ0XVHXPRI1DWXUDO+LVWRU\ biogeographic relationships of the mouse opossum THOMAS, O. 2QMicoureus griseus'HVPZLWK Thylamys (Didelphimorphia, Didelphidae) in description of a new genus and species of VRXWKHUQ6RXWK$PHULFD0ROHFXODU3K\ORJHQHWLFV 'LGHOSKLGDH$QQDOVDQG0DJD]LQHRI1DWXUDO DQG(YROXWLRQ +LVWRU\ PATTON, J.L., DA SILVA, M.N.F. & MALCOLM, J.R THOMAS,O. 2QQHZVPDOOPDPPDOVIURPWKH Gene genealogy and differentiation among 1HRWURSLFDO5HJLRQ$QQDOVDQG0DJD]LQHRI arboreal spiny rats (Rodentia, Echimyidae) of 1DWXUDO+LVWRU\ the Amazon basin - a test of the riverine barrier THOMAS,O. 2QMarmosa marmota and elegans, K\SRWKHVLV(YROXWLRQ with descriptions of new subspecies of the PATTON, J.L., DOS REIS, S.F. & DA SILVA, M.N.F. ODWWHU$QQDOVDQG0DJD]LQHRI1DWXUDO+LVWRU\ Relationships among didelphid marsupials based on sequence variation in the mitochondrial THOMAS,O. 7KUHHVPDOOPDPPDOVIURP6RXWK F\WRFKURPHEJHQH-RXUQDORI0DPPDOLDQ $PHULFD$QQDOVDQG0DJD]LQHRI1DWXUDO+LVWRU\ (YROXWLRQ PATTON, J.L., DA SILVA, M.N.F. & MALCOLM, J.R. THOMAS,O. 7KUHHQHZVSHFLHVRIMarmosa with Mammals of the Rio Juruá and the evolutionary note on Didelphys waterhousei 7RPHV$QQDOVDQG DQGHFRORJLFDOGLYHUVLILFDWLRQRI$PD]RQLD 0DJD]LQHRI1DWXUDO+LVWRU\ Bulletin of the American Museum of Natural THOMPSON, J.D., GIBSON, T.J., PLEWNIAK, F., JEANMOUGIN, History F. & HIGGINS, D.G., 7KH&/867$/;
197 Gayana 73(2), 2009
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