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Mus eum of Te x a s Te c h U n i v e r s i t y
Number 62
xxx
day
17month
June 2014
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Front cover: Collection localities and estimated ranges for 12 Bolivian species of Ctenomys: C. boliviensis,
C. conoveri, C. frater, C. leucodon, C. lewisi, C. nattereri, C. opimus, C. steinbachi, and the four new species
described herein, C. andersoni n. sp., C. erikacuellarae n. sp., C. lessai n. sp., and C. yatesi n. sp. Shown also
is a single locality for C. bicolor)LJXUHPRGL¿HGIURP*DUGQHU (1991).
6ƠƕƓƙƑƜ3ƥƒƜƙƓƑƤƙƟƞƣ
Museum of Texas Tech University
!Number 62
New Species of Ctenomys Blainville 1826 (Rodentia:
Ctenomyidae) from the Lowlands and Central Valleys
of Bolivia
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Layout and Design:
&RYHU'HVLJQ Production Editor:
Lisa Bradley
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Lisa Bradley
Copyright 2014, Museum of Texas Tech University
This publication is available free of charge in PDF format from the website of the Natural Science
Research Laboratory, Museum of Texas Tech University (nsrl.ttu.edu). The authors and the Museum
of Texas Tech University hereby grant permission to interested parties to download or print this
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other works is not permitted without prior written permission of the Museum of Texas Tech University.
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the Council on Library Resources.
Printed:
17 June 2014
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Special Publications, Museum of Texas Tech University, Number 62
Series Editor:
Robert J. Baker
New Species of Ctenomys Blainville 1826 (Rodentia: Ctenomyidae) from the Lowlands and
Central Valleys of Bolivia
6FRWW/*DUGQHU-RUJH6DOD]DU%UDYRDQG-RVHSK$&RRN
,661
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,6%1
Museum of Texas Tech University
/XEERFN7;86$
!"#$%&"'("%$)*$!"#$%&'($+,-(!.(,,"$/012$34)5"!6(-7$'6"!)89(5-":$*4)8$
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-+%64-'6
The genus Ctenomys%ODLQYLOOHLVRQHRIWKHPRVWGLYHUVHRI6RXWK$PHULFDQK\VWUL
FRJQDWKURGHQWV&XUUHQWO\QLQHVSHFLHVRIWXFRWXFRVDUHUHSRUWHGIURP%ROLYLDIRXUDWHOHYDWLRQV
DERYHPDQG¿YHLQKDELWLQJWKHORZODQGVP,QWKHSUHVHQWSDSHUPRUSKRORJ\
NDU\RORJ\DQGSK\ORJHQHWLFDQDO\VHVRI'1$VHTXHQFHVIRUDPLWRFKRQGULDOORFXVZHUHXVHG
to assess the taxonomic status of specimens of Ctenomys from localities beyond the previously
NQRZQUDQJHVRIWKHVHURGHQWVLQWKHGHSDUWPHQWVRI&KXTXLVDFD&RFKDEDPED6DQWD&UX]
and Tarija. Based on these analyses, we describe four new species in the genus Ctenomys, all
apparently endemic to the country. In addition, we place Ctenomys goodfellowi Thomas 1921
in synonymy under C. boliviensis:DWHUKRXVHDQGFRQ¿UPWKHSUHVHQFHRIC. nattereri
:DJQHUDVDGHQL]HQRIWKHHDVWHUQORZODQGVWKHUHIRUHWKHWRWDOQXPEHURIGRFXPHQWHG
extant species of Ctenomys in Bolivia is now 12.
Key words: Bolivia, CtenomysF\WRFKURPHbLQWHU$QGHDQYDOOH\VVXEWHUUDQHDQURGHQWV
WXFRWXFR<XQJDV
4"%<8"!
El género Ctenomys Blainville 1826 es uno de los géneros de roedores histricognatos más
GLYHUVRVGH$PpULFDGHO6XU1XHYHHVSHFLHVGHWXFRWXFRVHKDQUHSRUWDGRHQ%ROLYLDFXDWUR
HQHOHYDFLRQHVPD\RUHVDP\FLQFRTXHKDELWDQHQODVOODQXUDWURSLFDOHV\FKDTXHxDV
P(QHOSUHVHQWHWUDEDMRXWLOL]DPRVFDUDFWHUHVPRUIROyJLFRV\FDULROyJLFRVORVTXH
FRPELQDGRVFRQORVUHVXOWDGRVGHDQiOLVLV¿ORJHQpWLFRVGHPDUFDGRUHVPROHFXODUHVVHFXHQ
FLDVGH$'1SDUDXQORFXVPLWRFRQGULDOQRVVLUYLHURQSDUDHYDOXDUHOHVWDWXVWD[RQyPLFRGH
especímenes de Ctenomys SURYHQLHQWHVGHORFDOLGDGHVHQORVGHSDUWDPHQWRVGH&KXTXLVDFD
&RFKDEDPED6DQWD&UX]\7DULMD\TXHKDVWDHOPRPHQWRQRKDEtDQVLGRDQDOL]DGRVHQGHWDOOH
Sobre la base de estos análisis, se describen cuatro especies nuevas en el género Ctenomys,
WRGDVDSDUHQWHPHQWHHQGpPLFDVGHOSDtV$GHPiVHVWRVDQiOLVLVQRVSHUPLWHQVLQRQLPL]DU
Ctenomys goodfellowi bajo C. boliviensis:DWHUKRXVH\FRQ¿UPDUODSUHVHQFLDGHC.
nattereri:DJQHUFRPRXQKDELWDQWHGHODVWLHUUDVEDMDVGHORULHQWHSRUORWDQWRHQHVWH
momento existen 12 especies documentadas de Ctenomys en Bolivia.
3DODEUDV FODYHV %ROLYLD FLWRFURPRb, Ctenomys URHGRUHV VXEWHUUiQHRV WXFRWXFR
9DOOHV,QWHUDQGLQRV<XQJDV
Online supplemental documentation available at http://digitalcommons.unl.edu/parasitologyfacpubs/722.
1
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Ctenomys Blainville 1826 (Rodentia: Ctenomyidae).
7KHVHURGHQWVDUHHQGHPLFWR6RXWK$PHULFDDQGRF
FXU LQ ZHOOGUDLQHG IULDEOH VRLOV IURP DERXW ODWLWXGH
12°S in southern Peru (Sanborn and Pearson 1947),
south to Tierra del Fuego, and from the highlands of
3HUXDQG%ROLYLDWRWKHORZODQGVRIVRXWKHDVWHUQ%UD]LO
DQGQRUWKHDVWHUQ8UXJXD\3HDUVRQDQG&KULVWLH
:RRGDQG.LOSDWULFN&XHWRHWDO0DQ\
aspects of the biology of ctenomyids remain unstudied,
and in many cases distributional limits of species are
SRRUO\GH¿QHG(VWLPDWHVRIWKHQXPEHURIVSHFLHVLQ
WKHJHQXVUDQJHIURPWRPRUHWKDQ:RRGVDQG
Kilpatrick 2005).
Early studies on the systematics and taxonomy
of Ctenomys relied on color of pelage and general
morphological analyses of skull and other characters
RIWKHERG\/DWHUFKURPRVRPHVDOOR]\PHVDQG'1$
VHTXHQFHVZHUHXVHGWRH[DPLQHWKHV\VWHPDWLFVDQG
ELRJHRJUDSK\RIWKHVHURGHQWV&RRN*DUGQHU
&RRN DQG<DWHV /HVVD DQG &RRN &RRNDQG/HVVD0DVFKHUHWWLHWDO&RRN
DQG6DOD]DU%UDYR&DVWLOORHWDO)UHLWDV
3DUDGDHWDO)UHLWDVHWDO
In lowland Bolivia, these rodents reach a north
ernmost limit of about 16°S in the department of Santa
&UX]$WKLJKHUHOHYDWLRQVLQPRXQWDLQVDQGRQWKH
DOWLSODQRRIZHVWHUQ%ROLYLDWXFRWXFRVRFFXUVRXWKRI
/DNH7LWLFDFDFDP$QGHUVRQHWDO
showed that four species occur in the eastern lowlands
of Bolivia including C. boliviensis Waterhouse 1848,
C. steinbachi Thomas 1907 (see Thomas 1907, Wagner
8
1948,
and Waterhouse 1848), C. conoveri Osgood 1946
(see Osgood 1946), and a species that was referred
to as C. minutus Nehring 1887. Cook et al. (1990)
showed that four species of Ctenomys occurred at high
elevations (> 2,000 m) including: Ctenomys opimus
Wagner 1848, C. leucodon Waterhouse 1848, C. lewisi
Thomas 1926, and C. frater Thomas 1902 (see Thomas
1902, 1926). Ctenomys goodfellowi Thomas 1921
was considered a species different from C. boliviensis
E\&RRNDQG<DWHVDFRQFOXVLRQDFFHSWHGE\
$QGHUVRQ
Ctenomyids occupying habitats in intermediate
HOHYDWLRQVLQFOXGLQJWKH$QGHDQYDOOH\VRIFHQWUDO%R
OLYLDKDYHQRWEHHQVWXGLHGLQGHWDLO$QGHUVRQ
In the present paper, we examined all species of CtenomysLQDQGDURXQG%ROLYLD¿UVWXVLQJPLWRFKRQGULDO
'1$VHTXHQFHVWRGH¿QHVSHFLHVOLPLWVDQGUHODWLRQ
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morphological, and karyological variation.
Species of Ctenomys have mostly allopatric
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in morphological and chromosomal characters. To
identify species, we used the principles of concordance
$YLVHDQGUHFLSURFDOPRQRSK\O\7KHUHIRUHZH
UHFRJQL]H VSHFLHV DV WKH PRVW H[FOXVLYH KLHUDUFKLFDO
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diagnosable and reciprocally monophyletic lineages.
8-6"4(-,%$-!5$8"6;)5%
Field collecting, specimens, and cytogenetics.—
$QLPDOVZHUHFROOHFWHGXVLQJWUDSVVHWLQEXUURZVIRO
ORZLQJJXLGHOLQHVDSSURYHGE\WKH$PHULFDQ6RFLHW\
of Mammalogists (Sikes et al. 2011). Individuals were
measured, weighed, determined to sex, and prepared
DVPXVHXPVSHFLPHQV<DWHVHWDOZLWKWLVVXH
VDPSOHVDQGNDU\RW\SHVIUR]HQLQOLTXLGQLWURJHQ&RRN
$OO DUWKURSRG DQG KHOPLQWK SDUDVLWHV ZHUH
DUFKLYHGIROORZLQJVWDQGDUGSURWRFROVHJ*DUGQHU
*DUGQHUDQG-LPpQH]5XL]5HSUHVHQWD
tive individuals from most populations were karyotyped
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%DNHU HW DO DV PRGL¿HG E\$QGHUVRQ HW DO
$WOHDVWPHWDSKDVHFHOOVZHUHSKRWRJUDSKHG
and scored to determine diploid and fundamental num
bers. Fundamental number (FN) of chromosomal arms
IROORZVWKH³DXWRVRPDOQXPEHU´FRQYHQWLRQ*DUGQHU
and Patton 1976). For each individual specimen
VWXGLHGIXUDQGWHHWKFRORUZHUHFKDUDFWHUL]HGXQGHU
natural light. For both descriptions and comparisons
of species, we followed the color nomenclature of
Ridgway (1912).
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Specimens examined are archived in these mu
seum collections: Division of Mammals, Museum
RI6RXWKZHVWHUQ%LRORJ\06%$OEXTXHUTXH1HZ
0H[LFR86$'HSDUWPHQWRI0DPPDORJ\$PHULFDQ
0XVHXPRI1DWXUDO+LVWRU\$01+1HZ<RUN&LW\
&ROHFFLyQ%ROLYLDQDGH)DXQD&%)/D3D]%ROLYLD
and Museo “Noel Kempff Mercado” (MNKM), Santa
&UX]%ROLYLD)UR]HQWLVVXHVFHOOVXVSHQVLRQVDQG
slides with stained chromosome spreads are deposited
LQWKH'LYLVLRQRI*HQRPLF5HVRXUFHV'*5RIWKH
06%$Q\SDUDVLWHVFROOHFWHGGXULQJRXU¿HOGH[SH
ditions are archived in the Bolivian Mammal Parasite
Collection in the Harold W. Manter Laboratory of
Parasitology (HWML), University of Nebraska State
0XVHXP/LQFROQ1HEUDVND86$
Standard external measurements recorded for
each individual at time of collection include: TL:
WRWDOERG\OHQJWK7WDLOOHQJWK+)KLQGIRRWOHQJWK
LQFOXGLQJWKHFODZ(/HDUOHQJWKDQG:ZHLJKW
6NXOO PHDVXUHPHQWV DQG FKDUDFWHU GH¿QLWLRQV )LJ
IROORZ$QGHUVRQHWDODQG*DUGQHU
and included: condylobasilar root length (CBR),
condyloincisive length (CIL), palatal root length
(PLR), alveolar molar length (MOL), palatal breadth,
PLQLPXP3$%GHQWDOVSDQEUHDGWK'6%]\JRPDWLF
EUHDGWK=<%ODPEGRLGDOEUHDGWK/$%LQWHURUELWDO
EUHDGWK,2%DQGKHLJKWRIVNXOO6+7$GGLWLRQDO
TXDOLWDWLYHFKDUDFWHUVLQFOXGHGFXUYDWXUHRI]\JRPDWLF
SURFHVV&=3MXJDOSURFHVVRI]\JRPDWLFDUFK3=$
incisive foramen (IF), frontal parietal foramen (FPF),
DUWLFXODWLRQ VXUIDFH RI FRQG\ORLG SURFHVV $6 DQG
occipital crest (OC). Nomenclature used to describe
VNXOOFKDUDFWHUVIROORZV/DQJJXWKDQG$EHOOD
and Wahlert (1974, 1985).
Taxonomic names follow Woods and Kilpatrick
XQOHVV PRGL¿HG E\ PRUH UHFHQW WD[RQRPLF
FKDQJHV HJ )UHLWDV HW DO 6WRO] HW DO ,QIRUPDO QDPHV DVVRFLDWHG ZLWK VSHFLPHQV DQDO\]HG
by previous authors are included in the synonymies.
Morphological and morphometric analyses.—
Morphological comparisons and morphometric
analyses of the Ctenomys species were guided by
phylogenetic analyses of molecular data (see below).
Specimens were allocated into three age groups fol
ORZLQJ /DQJJXWK DQG$EHOOD DQG$OWXQD DQG
Lessa (1985). Pooled (male and female) measurements
B
of skull, weight, and external dimensions are given in
Table 1. Measurements of all individuals included in
WKLV VWXG\ DUH JLYHQ IRU HDFK VSHFLHV LQ 7DEOH6 LQ
online supplemental documentation.
6$6VRIWZDUHZDVXVHGIRUDOOPXOWLYDULDWH
VWDWLVWLFDODQDO\VHV%HFDXVHWKHIUHTXHQF\GLVWULEX
tions of some characters deviated slightly from nor
mality, all data were log transformed (log10) prior to
analysis. For the morphometric variables listed above,
stepwise discriminant function analysis (STEPDISC)
ZDVHPSOR\HGWRH[SORUHWKHGDWDVHWDQGDWWHPSWWR
identify diagnostic mensural characters. Canonical
GLVFULPLQDQWIXQFWLRQDQDO\VLV&$1',6&ZDVDOVR
used to establish a linear combination of morphological
YDULDEOHVWKDWEHVWGHOLQHDWHGVSHFLHVWKDWZHUHGH¿QHG
a priori on the basis of morphological comparisons,
chromosomes, and phylogenetic analyses. These vari
ables were then used to highlight the similarities and
differences between and among species of Ctenomys
that we studied.
0HWKRGVIRU'1$H[WUDFWLRQDPSOL¿FDWLRQDQG
VHTXHQFLQJZHUHDVJLYHQLQ/HVVDDQG&RRN
Parada et al. (2011), and Freitas et al. (2012). Each
3&5IUDJPHQWZDVVHTXHQFHGLQERWKGLUHFWLRQVVH
TXHQFHVZHUHIUHHRILQGHOVSUHPDWXUHVWRSFRGRQV
and ambiguities in forward and reverse directions,
providing support for a mitochondrial origin (Triant and
'H:RRG\6HTXHQFHGDWDWKDWZHJHQHUDWHGIRU
¿YHLQGLYLGXDOVLQFOXGHGLQWKLVSDSHUKDYHEHHQGHSRV
LWHGLQ*HQ%DQN$OOVSHFLHVQDPHVXVHGLQRXUVWXG\
DORQJZLWKVSHFL¿F¿HOGFDWDORJPXVHXPFDWDORJDQG
RU*HQ%DQNDFFHVVLRQQXPEHUVDUHJLYHQLQ7DEOH6
in the online supplemental documents.
DNA sequence analyses.²*HQHWLFFRPSDULVRQV
and phylogenetic analyses were based on complete cy
WRFKURPHb gene (cytEVHTXHQFHVFRQVLVWLQJRI
EDVHSDLUVESZLWKWKHH[FHSWLRQRIVHTXHQFHVHJ
C. bicolor) which ranged in length from 806 to 1,127
nucleotides. Our dataset differs from that of Freitas et
al. (2012), the most inclusive analysis of cytEVHTXHQFH
GDWDWRGDWHLQWKDWZHLQFOXGHG¿YHDGGLWLRQDO%ROLY
ian samples and representatives of both C. bicolor
and C. nattereriIURP%UD]LO,QWKHSUHVHQWSDSHUZH
generated a matrix of 75 terminal taxa representing
DSSUR[LPDWHO\FXUUHQWO\UHFRJQL]HGVSHFLHV
C$
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Figure 1. Dimensions of the cranium and mandible used in the descriptions and morphometric analyses of species
of CtenomysLQFOXGHGLQWKLVVWXG\IRUDGGLWLRQDOGH¿QLWLRQVVHH*DUGQHUDQG*DUGQHUDQG$QGHUVRQ
Measurements include: condylobasilar root length (CBR), condyloincisive length (CIL), palatal root length (PLR),
DOYHRODUPRODUOHQJWK02/SDODWDOEUHDGWKPLQLPXP3$%GHQWDOVSDQEUHDGWK'6%]\JRPDWLFEUHDGWK=<%
ODPEGRLGDOEUHDGWK/$%LQWHURUELWDOEUHDGWK,2%DQGKHLJKWRIVNXOO6+7$GGLWLRQDOTXDOLWDWLYHFKDUDFWHUVWKDW
ZHUHVWXGLHGLQFOXGHGFXUYDWXUHRI]\JRPDWLFSURFHVV&=3MXJDOSURFHVVRI]\JRPDWLFDUFK3=$LQFLVLYHIRUDPHQ
,)IURQWDOSDULHWDOIRUDPHQ)3)DUWLFXODWLRQVXUIDFHRIFRQG\ORLGSURFHVV$6RFFLSLWDOFUHVW2&/DEHOVRQ¿JXUH
DUH$YHQWUDOYLHZRIFUDQLXP%GRUVDOYLHZRIFUDQLXP&GRUVDOYLHZRIPDQGLEOH'OHIWODWHUDOYLHZRIVNXOO
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
D
Table 1. Descriptive statistics for external and cranial measurements for four new species of Ctenomys from
Bolivia. Measurements for sexes (see Materials and Methods text and Fig.1 for abbreviations) are pooled in
this table (see Table-S3 in online supplemental documentation for statistical summary and measurements of
individual specimens). All measurements are given as mean ± 1 standard deviation, minimum and maximum
measurements, and sample size (in parentheses).
Ctenomys
erikacuellarae n.sp.
Ctenomys yatesi
n.sp.
Ctenomys andersoni
n.sp.
Ctenomys lessai
n.sp.
TOTL
259.2 ± 28.2
±
209.5 ± 10.5
±
“
±
“
177 – 265 (9)
7$,/
72.9 ± 9.4
25 – 96 (68)
60.5 ± 2.5
±
69.8 ± 10
±
“
44 – 79 (9)
HFL
“
28 – 44 (69)
“
±
“
21 – 41 (27)
“
±
($5
7 ± 1.1
5 – 10 (69)
“
±
6.8 ± 1.4
±
6.6 ± 1.2
4 – 8 (9)
WT
222.1 ± 66.8
±
“
±
219.9 ± 60.8
±
175.8 ± 52
98 – 286 (9)
CBR
“
“
“
“
CIL
“
“
41.4 ± 4.4
±
“
PLR
“
15.2 ± 1
“
“
15.6 – 20 (6)
MOL
10.1 ± 0.8
8 ± 0.7
9.4 ± 0.7
9 ± 0.8
3$%
“
1.9 ± 0.1
±
“
“
DSB
9.8 ± 0.8
7.8 ± 0.6
9.5 ± 0.8
“
8 – 10 (6)
=<%
28.6 ± 2.6
21.8 ± 1.5
26.1 ± 2.5
26.4 ± 1.9
±
/$%
27 ± 2
22.1 ± 1.2
25.2 ± 2
“
IOB
11.2 ± 1
8.7 – 14.1 (56)
7.9 ± 0.6
9.4 ± 1
“
SHT
15.4 ± 1.4
“
±
“
“
Character
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Phylogenetic analysis.—Phylogenetic relation
ships based on cytE JHQH VHTXHQFHV ZHUH DVVHVVHG
XVLQJPD[LPXPSDUVLPRQ\03LQ3$836ZRIIRUG
$0D[LPXP/LNHOLKRRG0/WUHHZDVJHQHU
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3$83 ZDV XVHG WR FRQGXFW KHXULVWLF VHDUFKHV ZLWK
UHSOLFDWHVUDQGRPWD[RQDGGLWLRQ57$DQGWUHH
ELVHFWLRQUHFRQQHFWLRQEUDQFKVZDSSLQJ7%5ZLWK
DOOGDWDFRQVLGHUHGXQRUGHUHGDQGHTXDOO\ZHLJKWHG
1RQSDUDPHWULFERRWVWUDSDQDO\VHV)HOVHQVWHLQ
SVHXGRUHSOLFDWHVDQGUDQGRPVHTXHQFHDGGL
tions with each replicate) were run to assess support for
individual nodes. Nodes with bootstrap support above
ZHUHFRQVLGHUHGZHOOVXSSRUWHG0U0RGHO7HVW
1\ODQGHUZDVHPSOR\HGWR¿QGWKHHYROXWLRQDU\
PRGHOWKDWEHVW¿WWHGRXUcytEGDWDEDVHGRQWKH$NDLNH
LQIRUPDWLRQ FULWHULD WKH *HQHUDO 7LPH 5HYHUVLEOH
model of substitution, with a gamma distribution and
DSURSRUWLRQRI LQYDULDEOHVLWHV*75 * ,ZDV
GHWHUPLQHGWREHWKHEHVW¿WIRURXUcytb dataset and
ZDVWKHPRGHORIHYROXWLRQXVHGLQRXU5$[0/UXQV
2,500 ML rapid bootstrap analyses were run to estimate
the level of support for different clades in our result
ing trees. Pairwise genetic distances were calculated
to assess within and among species differences using
WKH ³S´ XQFRUUHFWHG GLVWDQFH HVWLPDWRU LQ 0(*$ (Tamura et al. 2011).
4"%<,6%
*HQHWLF GLVWDQFHV 7DEOH IRU WKH cytb gene
between and among Bolivian species of Ctenomys
averaged 8.8% and were slightly smaller when all
VSHFLHVZHUHLQFOXGHGVHH7DEOH6LQRQOLQH
supplemental documents). The smallest distance
among Bolivian species was 5% and the greatest was
7DEOHZKHQDOORWKHUVSHFLHVRICtenomys
ZHUHFRQVLGHUHG7DEOH6WKHVHHVWLPDWHVZHUH
and 12.8%, respectively.
cent common ancestor with C. conoveri and these two
VSHFLHVDUHSDUWRIDZHOOVXSSRUWHGFODGHWKDWDOVRLQ
cludes C. lewisi and C. frater (see descriptions below).
Results from phylogenetic analysis using both
maximum parsimony (MP) and maximum likelihood
(ML) as optimality criteria of our cytEGDWDVHWVKRZHG
congruent topologies relative to the Bolivian species
of Ctenomys. For the MP tree (Fig. 2) the pertinent
VWDWLVWLFVIRUFKDUDFWHUVZHUHWUHHOHQJWK FRQ
VLVWHQF\LQGH[LVUHWHQWLRQLQGH[LV$VLQ
previous studies (e.g., Parada et al. 2011, Freitas et al.
2012) our phylogenetic analysis of all available cytb
data also recovered various species groups with rela
WLYHO\KLJKOHYHOVRIVXSSRUWVHYHUDORIWKHVHVSHFLHV
and species groups occur in Bolivia. Our phylogenetic
K\SRWKHVHV )LJV DQG VKRZ WKUHH XQGHVFULEHG
%ROLYLDQVSHFLHV)LJ$DUURZ³D´WKDWFRPSULVHD
PRQRSK\OHWLFJURXS,QDGGLWLRQRXUDQDO\VHVFRQ¿UP
C. bicolor Miranda Ribeiro 1914 as the sister species to
C. nattereri Wagner 1848 which, as shown recently by
6WROW]HWDOLVXQHTXLYRFDOO\SUHVHQWLQ%ROLYLD
In addition, this tree (Fig. 2B, arrow “b”) indicates that
the undescribed species (NK22840) shares a most re
To assist in description and exploration of skull
morphology and to provide a basis for identifying
distinguishing characters of Ctenomys, we examined
PRUSKRPHWULFYDULDWLRQEDVHGRQDSUHYLRXVO\GH¿QHG
VHWRITXDQWLWDWLYHFKDUDFWHUV)RUWKHFRPSOHWHGDWD
set with 11 species from Bolivia, we investigated all
variables of the skull using discriminant analysis with
ERWKVWHSZLVH67(3',6&DQGFDQRQLFDO&$1',6&
analyses. We found that using the stepwise analysis,
the characters most important in discrimination among
JURXSV DUH 02/ 6+7 =<% 3/5 '6% 7DEOH Results from a standard canonical discriminant analy
VLVVKRZWKDW02/=<%'6%,2%DQG6+7EHVW
discriminate among species (see plot, Fig. 4, and the
IXOO6$6SORWRXWSXWLQWKH¿OHQDPHG³0XOWLY6WDWV
pdf” in online supplemental documents.). Therefore,
by combining results from both multivariate analyses,
common characters of the skull that enable best dis
crimination among all Bolivian species of Ctenomys
DSSHDUWRLQFOXGH02/=<%'6%DQG6+7
Like the MP analysis, the tree from the ML
DQDO\VLV)LJEDVHGRQXQLTXHKDSORW\SHVVKRZVD
topology congruent with previous studies, including the
short branches at the base of the radiation indicating
UDSLGGLYHUVL¿FDWLRQRIVSHFLHV/HVVDDQG&RRN
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
E
Table 2. Mean pairwise uncorrected cytochrome-b p-distances (scaled as percent sequence divergence) among Bolivian species of CtenomysEHORZGLDJRQDODQGLQWUDVSHFL¿FSGLVWDQFHVGLDJRQDO&RPSDULVRQVRIGLVWDQFHVRI
outgroup taxa with the ingroup Ctenomys sp. show much greater distances (>18%). A total of 1,140 positions in the
¿QDOF\WRFKURPHb dataset were included. See the complete matrix of genetic distances based on cytochrome-b in the
online supplemental material.
1
2
4
5
6
7
8
9
10
11
1
C. andersoni
n/c
2
C. boliviensis
6.4
1.2
C. conoveri
9.5
10.0
n/c
4
C. erikacuellarae
5.0
5.8
10.1
1.8
5
C. frater
10.4
10.9
9.2
1.2
6
C. lessai
10.5
10.7
6.8
10.4
8.6
n/c
7
C. leucodon
9.0
9.1
12.5
9.0
12.1
11.6
n/c
8
C. lewisi
10.4
10.5
8.2
10.7
11.8
n/c
9
C. nattereri
7.0
5.6
9.7
11.0
10.1
8.8
10.1
1.0
10
C. opimus
6.6
6.4
9.2
6.9
11.4
9.6
9.0
10.7
7.1
0.2
11
C. steinbachi
6.5
10.0
6.2
11.0
11.2
9.9
10.7
6.8
7.5
n/c
12
C. yatesi
5.4
6.5
9.2
5.7
10.2
10.0
9.5
9.9
6.8
6.9
12
0.2
0$
$$$$$$$$$)''-%()!-,$&-&"4%=$8<%"<8$)*$6">-%$6"';$<!(."4%(69
A)
a
)LJXUH5HVXOWVRIWKHPD[LPXPSDUVLPRQ\SK\ORJHQHWLFDQDO\VLVLQ3$83IRUWD[DRICtenomys
and three outgroup taxa that included the octodontids: Tympanoctomys barrerae (Lawrence 1941),
Octodon degus (Molina 1782), and Spalacopus cyanus0ROLQD$ERWWRPKDOIRIWKHWUHH
B) top half of the tree. See text for explanation of a and b arrows. The analysis was run on 1,140
XQRUGHUHGHTXDOO\ZHLJKWHGFKDUDFWHUVRIWKHFRPSOHWHPLWRFKRQGULDOF\WRFKURPHb (cytb) gene. The
tree depicted is the result of a heuristic search with 500 replicates of random taxon addition and TBR
DQG57$LQ3$831RQSDUDPHWULFERRWVWUDSDQDO\VLVZDVUXQWRDVVHVVEUDQFKVXSSRUWERRWVWUDS
values are mapped).
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
B)
b
Figure 2 (continued).
F
/G$
$$$$$$$$$)''-%()!-,$&-&"4%=$8<%"<8$)*$6">-%$6"';$<!(."4%(69
)LJXUH5HVXOWVRIDUXQZLWK5$[0/RIPD[LPXPOLNHOLKRRGUDSLGERRWVWUDSDQDO\VHVRIRXUctyb dataset. The
general topological groups seen in Figure 2 are evident in this tree. Numbers on the tree represent bootstrap values.
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
//
Table 3. Results of a stepwise discriminant analysis of log10 transformed variables for 11 species of Ctenomys from
Bolivia (including C. erikacuellarae n. sp. C. andersoni n. sp., C. lessai n. sp., C. yatesi n. sp., C. frater, C. boliviensis,
C. opimus, C. lewisi, C. conoveri, C. leucodon, and C. steinbachi) (individuals of C. nattereri not included here). The
most important characters discriminating among the species included in this analysis are listed. Those with the greatest
)YDOXHWKH¿UVWFKDUDFWHUVDSSHDUPRVWLPSRUWDQWLQWKHFDQRQLFDOGLVFULPLQDWLRQDPRQJVSHFLHV
Step
Character
Entered
3DUWLDO56TXDUH
F Value
Pr > F
Wilks’ Lambda
3U/DPEGD
1
MOL
80.17
2
SHT
0.2998
8.91
0.14478849
=<%
0.4662
18.08
0.07728911
4
PLR
0.2899
8.41
0.05488471
5
DSB
0.2128
5.54
6
CIL
0.1914
7
CBR
2.21
0.0184
8
/$%
0.0961
2.15
0.0225
9
3$%
0.0800
1.75
0.0726
0.02619546
Figure 4. Plot of canonical scores of mensural data of cranial measurements for 11 species of Ctenomys from Bolivia.
7KHDQDO\VLVZDVUXQXVLQJFDQRQLFDOGLVFULPLQDQWDQDO\VLV&$1',6&6$60LQLPXPSRO\JRQVUHSUHVHQWWKH
SORWVRILQGLYLGXDOFDQRQLFDOVFRUHVIRUD[HV,DQG,,7KH¿UVWWZRD[HVDFFRXQWIRURIWKHYDULDWLRQLQWKHGDWD
set. Zygomatic breadth and interorbital breadth contribute most to discrimination between and among species. See
RQOLQHVXSSOHPHQWDOGRFXPHQWDWLRQ0XOWLY6WDWVSGIIRUFRPSOHWHGDWDVHWDQGUHVXOWVRIWKH&$1',6&DQDO\VLV
/1$
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6->)!)89$-!5$%&"'("%$5"%'4(&6()!%
Based on the patterns of phylogenetic, karyotypic,
and morphological variation in specimens collected
from several localities in the valleys and lowlands
of Bolivia, four new species of Ctenomys (Rodentia
Bowdich 1821: Ctenomyidae Lesson 1842) are herein
described.
Ctenomys erikacuellarae, new species
(ULND¶V7XFR7XFR(O7XFR7XFRGH(ULND
Figs. 5, 6, 7, 8 (map), Table 1
CtenomysVSSDUW$QGHUVRQ
Ctenomys “monte”: Lessa and Cook (1998:89)
Ctenomys “mont”: Cook and Lessa (1998:1,524)
Ctenomys “Monte”: Castillo et al. (2005:172)
Ctenomys “monte”: Parada et al. (2011:676)
C. VS0217()UHLWDVHWDO
Holotype.²06%0XVHXPRI6RXWKZHVW
ern Biology), adult male. Skin, skull and axial skeleton
FROOHFWHG$XJXVWSUHSDUHGE\6\GQH\$QGHUVRQ
¿HOGQXPEHU6$$WWKHWLPHRIFROOHFWLRQWKH
OLYHU NLGQH\ DQG KHDUW WLVVXH ZHUH IUR]HQ LQ OLTXLG
QLWURJHQDQGVXEVHTXHQWO\VWRUHGDWƒ&LQWKH'LYL
VLRQRI*HQRPLF5HVRXUFHV0XVHXPRI6RXWKZHVWHUQ
Biology, NK21901.
Type locality.²%ROLYLD'HSDUWPHQWRI&KXTXL
VDFDNP(&KXKXD\DFRƒ
6ƒ
:HOHYDWLRQ
1,200 m.
Measurements of holotype.²727/7$,/
+)/ ($5 :7 &%5 &,/
3/5 02/ '6% 3$%
=<%,2%/$%6+7
Specimens examined.²6L[W\HLJKW DGGLWLRQDO
specimens from the type locality and two additional
ORFDOLWLHV LQ WKH 'HSDUWPHQWV RI 6DQWD &UX] DQG
&KXTXLVDFD %HVLGHV WKH KRORW\SH WKH UHVW RI WKH
series of specimens collected from the type local
LW\DUHGHVLJQDWHGSDUDW\SHVDVIROORZVQLQH$01+
±QLQH&%)
±DQGVL[06%
7KHUHVWRIWKHVSHFL
mens studied (but not designated as types) include the
IROORZLQJ7ZHQW\VHYHQLQGLYLGXDOVIURPNP6:
RI 0RQWHDJXGR P HOHYDWLRQ 'HSDUWPHQW RI
&KXTXLVDFD%ROLYLDƒ
6 ƒ
: $01+
±QLQH&%)
VHYHQ06%±
DQGWUDSSHGLQ-XO\
and 25 additional specimens, trapped in June 1991 from
near the Río Las Ciénegas, at 5.5 km NNE of Valle
JUDQGHPHOHYDWLRQ'HSDUWPHQWRI6DQWD&UX]
%ROLYLD ƒ
6 ƒ
: ¿YH 01. ±QLQH$01+06%
±±
67422, and 67424). Pooled measurements of skulls
DQGH[WHUQDOGLPHQVLRQVRIERWKVH[HVDUHVXPPDUL]HG
LQ7DEOHVHHDOVR7DEOH6LQRQOLQHVXSSOHPHQWDO
GRFXPHQWVIRUFRPSOHWHGDWDVHWDQGPHDVXUHPHQWVRI
individual specimens).
Diagnosis.²$ PHGLXPVL]HG WXFRWXFR RI WKH
“boliviensis” group of Ctenomys with dorsal and
YHQWUDOFRORUDWLRQZHOOGLIIHUHQWLDWHGGRUVXPPRVWO\
Ochraceous Orange except on the upper surface of
KHDG DQG PX]]OH ZKLFK LV EODFNLVK EURZQ YHQWHU
Drab Brown or Buffy Brown displaying extensive
white or Light Buff markings on inguinal, axillary
and/or pectoral regions. Skull strongly built, with
VWURQJO\FXUYHG]\JRPDWLFDUFKHVRUWKRGRQWLQFLVRUV
URVWUXPZLGHVWDWWKHWLSRISUHPD[LOODULHVDULEERQ
OLNH DOLVSKHQRLGSUHVSKHQRLG EULGJH HYHQ LQ \RXQJ
individuals, and a karyotype with 2N=24 and FN=40.
Description.²3HODJH GHQVH ¿QH VRIW DERXW
PPORQJRYHUEDFNDQGUXPSGRUVXPZLWKIXU
ranging from Ochraceous Orange to Buckthorn Brown,
KDLUVGDUNFRORUHGH[FHSWODVWRUPPZKLFKDUH
distinctly lighter. Most individuals with dark cap (Fus
cous Black) on head, ca. 1.5 to 2 cm wide, running from
MXVWDERYHQRVHWRDWOHDVWQHFNPLGGRUVDOVWULSHRI
same color running along back, and sometimes reach
ing rump. Color of ventral pelage more sharply set off
IURPGRUVDOLQVRPHLQGLYLGXDOVHJ$0WKDQ
LQRWKHUVHJ$0GHHS1HXWUDO*UD\EDVDOO\
ZLWKVXSHU¿FLDOWRPPZDVKRI'UDE%URZQRU
Buffy Brown. “Collar” of light fur extending ventrad
IURPSLQQDHWRJXODUUHJLRQHJ$0)XURI
fore and hind limbs colored like dorsum and rump,
except internal sides of both (axillary areas) which are
:KLWHWR/LJKW%XIILQVRPHLQGLYLGXDOVLQJXLQDODUHD
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
/B
)LJXUH9LHZVRIVNXOODQGPDQGLEOHRIKRORW\SH06%RICtenomys erikacuellaraeQVS$GRUVDOYLHZ
of skull, B) ventral view of skull, C) dorsal view of mandible, D) ventral view of mandible, E) en face view of skull
VKRZLQJGDUNRUDQJHFRORURIXSSHULQFLVRUV)OHIWODWHUDOYLHZRIVNXOO*OHIWODWHUDOYLHZRIPDQGLEOH
)LJXUH0XVHXPVWXG\VNLQRIKRORW\SH06%RICtenomys erikacuellaraeQVS$GRUVDOYLHZ%YHQWUDO
view, C) left lateral view.
/C$
$$$$$$$$$)''-%()!-,$&-&"4%=$8<%"<8$)*$6">-%$6"';$<!(."4%(69
)LJXUH6WDQGDUGJLHPVDVWDLQHGNDU\RW\SHSUHSDUDWLRQVRICtenomys erikacuellaraeQVS>1 )1 @
$KRORW\SHPDOH06%VKRZLQJPHWDFHQWULF;DQGDFURFHQWULF<FKURPRVRPHGLVWLQFWLYHRIWKLV
VSHFLHV%IHPDOHVKRZLQJWKHELDUPHGPHWDFHQWULF;;FKURPRVRPHV
FRORUHG:KLWHWR/LJKW%XIIHJ$0DQGLQ
others, with a pectoral shield of White or Light Buff
hairs. Top of feet covered with agouti hairs to base of
PHWDFDUSDOVDQGWKHQUHSODFHGE\ORQJHUVHOIFRORUHG
hairs. Mystacial vibrissae extending to the base of the
SLQQDH ZKHQ ODLG EDFN DORQJVLGH KHDG VXSHUFLOLDU\
vibrissae sparse, extending to the dorsal edge of the
SLQQDHZKHQODLGEDFNDORQJVLGHRIKHDGJHQDORQH
present, interramal present.
Ears sparsely covered with short, brownish
hairs, not contrasting conspicuously with color of
head. Opening to auditory canals protected by small
FWHQLIRUPVHOIFRORUHGEULVWOHVEXQGOHVRIVWLIIKDLUV
posterior to upper and lower incisors. Pes broad, all
digits with ungal tufts of stiff bristles (e.g. cteniform),
DQGVWURQJFODZV)RUHIRRWZLWKDZHOOGHYHORSHGSRO
lex bearing a short open nail, other four digits bearing
basally closed claws. Tail short, strong, teretiform
in cross section, darker above than below in most
specimens, sparsely covered with dark hairs, except
for terminal 5–7 mm, which are covered with whitish
hairs. Six mammae in inguinal, abdominal, and post
axial pairs, postaxial pair enlarged even in very young
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
)LJXUH0DSRI%ROLYLDPRGL¿HGIURP*DUGQHUZLWKFROOHFWLRQORFDOLWLHVRICtenomys species that we
FROOHFWHG IURP WR &RQWLQXRXVO\ VKDGHG DUHD WKURXJK WKH VRXWKHUQ KDOI RI %ROLYLD DQG LQWR %UD]LO
indicates approximate range of rodents of the genus Ctenomys. Included are approximate collection localities and
estimated ranges in Bolivia for C. opimus, C. leucodon, C. lessai n. sp., C. lewisi, C. frater, C. erikacuellarae n.
sp., C. steinbachi, C. boliviensis, C. yatesi n. sp., C. conoveri, and C. andersoni n. sp. Shown also are localities
and estimated range for C. nattereriIURP%ROLYLDDQG%UD]LOVHHWH[WDQGIRUDVLQJOHORFDOLW\IRUC. bicolor (skull
VSHFLPHQIURP$01+WKDWZDVH[DPLQHGE\XV
/D
/2$
$$$$$$$$$)''-%()!-,$&-&"4%=$8<%"<8$)*$6">-%$6"';$<!(."4%(69
IHPDOHV*ODQVSHQLVWRWLPHVORQJHUWKDQZLGHDQG
cylindrical, externally covered with coarse spines (well
developed in adults only).
6NXOO UREXVW LQWHURUELWDO UHJLRQ ZLWK VTXDUHG
PDUJLQV ]\JRPDWLF DUFKHV EURDG DXGLWRU\ EXOODH
LQÀDWHGS\ULIRPDXGLWRU\WXEHVVDOLHQW1DVDOERQHV
EURDGHVWDQWHULRUO\ZLWKÀDWGRUVDOHGJHVQDVDOVVKRUW
premaxillary clearly visible when viewed from dorsal
DVSHFW)URQWDOVZLGHVWDWOHYHORIFRQQHFWLRQRI]\
gomatic arches, lateral to posterior anterior sutures of
frontals, nasals, and maxillaries. Interorbital processes
of frontals almost as wide as widest part of frontals,
frontals constricting to their narrowest at level of
VXWXUHV RI IURQWDOV DQG SDULHWDOV SDULHWDOV QDUURZHVW
DWVLWHRIDWWDFKPHQWRIVTXDPRVDOURRWRI]\JRPDWLF
DUFK6TXDPRVDODSSHDULQJWRH[SDQGZLWKDJHDQGWKXV
in young individuals parietal and back of frontals are
about same width. In older individuals, interparietal
completely fused or extremely small and indistinct.
7HPSRUDOIRUDPHQDWMXQFWLRQRIVTXDPRVDODQGSDULHWDO
always present and not elongated. Lambdoidal ridge
always present, even in young individuals. Supraoc
cipital crest strongly developed in adults. Zygomatic
arch strongly built, with massive processes extending
ERWKGRUVDGDQGYHQWUDGMXJDOH[WHQGHGGRUVRYHQWUDO
O\LQFLVLYHIRUDPLQDUHFHVVHGLQDFRPPRQIRVVDDQG
incompletely separated by a bony septum (or if pres
HQWVHSWXPLVZHDNO\GHYHORSHGLQPRVWLQGLYLGXDOV
LQWHUSUHPD[LOODU\IRUDPHQFRQVSLFXRXVSDODWDOEULGJH
with two major palatine foramina at about level of M1.
Mesopterygoid fossae, an inverted “V” shape, reaching
anteriorly to level of M2. Posteropalatal pits present,
EXWQRWHQODUJHG$OLVSKHQRLGSUHVSKHQRLGEULGJHÀDW
DQG ULEERQOLNH HYHQ LQ \RXQJHU LQGLYLGXDOV %RQ\
roof of mesopterygoid fossa usually completely ossi
¿HGVRPHWLPHVZLWKWLQ\SHUIRUDWLRQVEXWQHYHUODUJH
VSKHQRSDODWLQH YDFXLWLHV EXFFLQDWRUPDVWLFDWRU\
foramen enlarged and undivided in many individuals
(including holotype), but divided into two or sometimes
three small foramina in other individuals. Upper inci
sors large, robust, Orange in color, and opisthodont.
Maxillary tooth rows slightly divergent posteriad.
Mandible robust, with coronoid process fal
FLIRUP QRW VWURQJO\ DQJOHG EDFNZDUGV FRQG\ORLG
SURFHVVVWURQJEHDULQJDQDUWLFXODWLRQÀDQJHQRWZHOO
developed.
Karyology.—2N=24 and FN=40. Karyotypes
were examined from seven individuals from 2 km
6: RI 0RQWHDJXGR RQH PDOH 06% DQG VL[
IHPDOHV06%$01+±&%)
1100–1101, 2654), six specimens from 2 km E of
&KXKXD\DFRRQHPDOH06%DQG¿YHIHPDOHV
$01+±&%)DQG
eight animals from 5.5 km NNE of Vallegrande (one
male and seven females). Typical karyotype (Fig.
VKRZLQJ QLQH SDLUV RI ELDUPHG DXWRVRPHV DQG
two pairs of acrocentric autosomes of which one is
PHGLXPVL]HGDQGGLVWLQFWLYHDQGWKHRWKHULVDSDLU
that is extremely small. Of the biarmed autosomes,
six pairs are submetacentric, and three pairs are
metacentric, large acrocentric pair, distinctive. The
; FKURPRVRPH LV ODUJH DQG PHWDFHQWULF WKH <
FKURPRVRPHLVDFURFHQWULFDQGDERXWKDOIWKHVL]HRI
WKH;FKURPRVRPH)LJ
Distribution and habitat.²(ULND¶V WXFRWXFR
is known only from three localities (Fig. 8) situated
RQWKHHDVWHUQÀDQNVRIWKH&RUGLOOHUD2ULHQWDORIWKH
$QGHV EHWZHHQ DQG PHWHUV DOWLWXGH RI
VRXWKFHQWUDO%ROLYLDLQWKH'HSDUWPHQWVRI&KXTXLVDFD
DQG6DQWD&UX]$PLQLPXPJHRJUDSKLFDUHDSRO\JRQ
connecting all three localities encompasses an area
VOLJKWO\ODUJHUWKDQNPðVHH)LJ)LQRQOLQH
supplemental documents).
$OOWKUHHNQRZQORFDOLWLHVDUHSDUWRIWKHHFRORJL
FDO]RQHNQRZQDV%RVTXHV6HFRV,QWHUDQGLQRV,ELVFK
HWDORU$QGHDQGU\YDOOH\V/RSH]7KH
locality designated “2 km SW of Monteagudo” was on
an agricultural experimental station. Vegetation was
less disturbed inside the fenced station and was typi
FDORIWKHORZHUHDVWHUQHVFDUSPHQWVRIWKH$QGHVLQ
VRXWKHUQ%ROLYLD,ELVFKHWDO
$WWKHFROOHFWLRQORFDOLW\³NP11(RI9DO
legrande” on the Río Ciénega, the hillsides were cov
HUHGZLWKPHVTXLWHProsopis L.), columnar cacti (e.g.
Trichocereus sp.), acacia (Acacia sp.), and Ximenia sp..
The Río Ciénega runs through the valley and is sur
rounded by dense stands of shrubs and trees (Prosopis,
HWF7KHDUHDZDVFRQYHUWHGWRFURSODQGDQGLVJUD]HG
heavily by livestock. The collecting locality at “2 km
East of Chuhuayaco” was similar to that of Vallegrande.
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
Individual tucos from Vallegrande have a distinct
FRORUDWLRQVRPHVKRZZKLWHPDUNLQJVEHKLQGWKHHDUV
and all had frontoparietal fontanelles or fenestrae. Else
where, the percentage of animals with fenestrae ranges
from 78% near Monteagudo to 92% in Chuhuayaco
*DUGQHUDQG$QGHUVRQ
Etymology.—We name this species in honor of
our friend and colleague Erika Cuéllar, a Bolivian
conservation biologist who has worked tirelessly to
SURWHFWWKHÀRUDDQGIDXQDRIWKH*UDQ&KDFRUHJLRQ
RI6RXWK$PHULFDE\HPSRZHULQJWKHSHRSOHLQORFDO
communities. In addition, Erika participated in the
¿HOGH[SHGLWLRQVWKDWVHFXUHGVRPHRIWKHHDUOLHVWVSHFL
PHQVRIWKLVSRRUO\NQRZQVSHFLHVRIWXFRWXFRZH
fondly remember her enthusiasm, positive attitude, and
LQTXLVLWLYHPLQGDQGKDLOWKHVHDVH[DPSOHVWRIROORZ
The species name is a patronym in the genitive singular.
Comparisons.—Ctenomys erikacuellarae n. sp.
FDQEHUHFRJQL]HGDVGLVWLQFWIURPVSHFLHVRICtenomys
known from in and around Bolivia by the following:
Ctenomys erikacuellarae differs from C. steinbachi, a
species occurring in lowland Bolivia near the city of
6DQWD&UX]GHOD6LHUUDE\KDYLQJQRFROODURIOLJKWFRO
ored fur, by having fur that is lighter and more Ochra
ceous in color and by having a diploid fundamental
chromosome number of 2N=24, FN=40 versus 2N=10,
FN=16 in C. steinbachi VHH$QGHUVRQHWDO
Ctenomys erikacuellarae is smaller than C.
conoveri and has fewer chromosomes (2N=48, FN=70
in C. conoveriVHH$QGHUVRQHWDODQGDVPRRWK
pelage relative to the shaggy coat of C. conoveri.
Ctenomys erikacuellarae can be differenti
ated from C. boliviensis by lacking a collar through
the gular region, a karyotype of 2N=24 and FN=40
versus 2N=42–46 and FN=64 in known populations
of C. boliviensisVHH$QGHUVRQHWDO smaller
RYHUDOO VNXOO VL]HV RI DGXOWV DQG KLJK SUHYDOHQFH RI
fenestrae at the frontal parietal suture versus very low
prevalence to none in C. boliviensisVHH*DUGQHUDQG
$QGHUVRQ
Ctenomys erikacuellarae can be distinguished
from C. frater by having rough, bicolored, Ochraceous
fur dorsally, while C. frater has fur that is very soft
/E
DQG6ODWH*UD\LQFRORUDQGDPXFKGLIIHUHQWORZHU
chromosome number versus a 2N=48 and FN=90 in C.
frater (see Cook et al. 1990).
Finally, C. erikacuellarae can be differentiated
IURP /HZLV¶ WXFRWXFR C. lewisi) by having Ochra
ceous fur (versus very dark fur in C. lewisi) and a
different karyotype (C. lewisi KDVD1 )1 Cook et al. 1990). In dimensions of the skull, C. erikacuellarae differs from C. lewisiLQPHDQVNXOOOHQJWK
FRQG\ORLQFLVLYHOHQJWKLQLQWHURUELWDOEUHDGWKDQG
mean skull height (Table 1).
Ctenomys yatesi, new species
<DWHV¶7XFR7XFRHOWXTXLWRGH<DWHV<DWHV¶WXTXLWR
Figs. 8 (map), 9, 10, Table 1
Ctenomys minimus$QGHUVRQPLVVSHOOLQJ
of minutus)
Ctenomys minutus: Olds et al. (1987:17)
Ctenomys minutus$QGHUVRQHWDO
Ctenomys minutus$QGHUVRQ
Ctenomys sp. (“minut”): Lessa and Cook (1998:89)
Ctenomys “min” : Cook and Lessa (1998:1,524)
Ctenomys sp. (“minut”): Slamovits et al.
(2001:1,709)
Ctenomys minutus*DUGQHUDQG$QGHUVRQ
C. sp. “Minut”: Castillo et al. (2005:172)
Ctenomys sp. MINUT: Parada et al. (2011:682)
CVS0,187)UHLWDVHWDO
Holotype.²$01+$PHULFDQ0XVHXP
of Natural History), adult male. Skin, skull, and axial
skeleton collected 9 October 1984, prepared by Sydney
$QGHUVRQ¿HOGQXPEHU6$$WWKHWLPHRIFROOHF
WLRQWLVVXHVRIWKHOLYHUNLGQH\DQGKHDUWZHUHIUR]HQ
LQ OLTXLG QLWURJHQ DQG VXEVHTXHQWO\ VWRUHG DW °C
LQ WKH 'LYLVLRQ RI *HQRPLF 5HVRXUFHV 0XVHXP RI
Southwestern Biology, NK12406.
Type locality.—Bolivia, Department of Santa
&UX] NP 1 DQG NP : RI 5RERUp ƒ
6
ƒ
:PHOHYDWLRQ
Measurements of holotype.²727/7$,/
+)/($5:7&%5&,/
3/502/'6%3$%=<%
,2%/$%6+7
/0$
$$$$$$$$$)''-%()!-,$&-&"4%=$8<%"<8$)*$6">-%$6"';$<!(."4%(69
)LJXUH9LHZVRIVNXOODQGPDQGLEOHRIKRORW\SH$0RICtenomys yatesiQVS$GRUVDOYLHZRI
skull, B) ventral view of skull, C) dorsal view of mandible, D) ventral view of mandible, E) en face view of
VNXOOVKRZLQJGDUNRUDQJHFRORURIXSSHULQFLVRUVDQGODUJHW\PSDQLFEXOODH)OHIWODWHUDOYLHZRIVNXOO*OHIW
lateral view of mandible.
)LJXUH0XVHXPVWXG\VNLQRISDUDW\SH&%)RICtenomys yatesiQVS$GRUVDOYLHZ%
ventral view, C) left lateral view.
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
Specimens examined.—Two additional speci
mens from the type locality. Designated as paratypes:
06%&%)3RROHGPDOHDQGIHPDOHPHD
surements of skulls, weights, and external dimensions
are given in Table 1. Individual measurements used
DUHVXPPDUL]HGIRUHDFKVSHFLHVLQ7DEOH6LQRQOLQH
supplemental documents.
Diagnosis.²$VPDOOVL]HGVSHFLHVRICtenomys,
member of the “boliviensis” group of this genus, dor
sally Pale Brown, even more pale ventrally, without a
hint of dark dorsal stripe and no markings on fur in the
JXODUUHJLRQYHU\ODUJHW\PSDQLFEXOODH)LJ(DQG
very distinctive range of morphometric variation (from
50 to 70% smaller than any other species of Ctenomys
in Bolivia).
Description.²3HODJH¿QHDQGVRIWKDLUVDERXW
PPORQJRYHUEDFNDQGUXPSGRUVDOFRORUDWLRQ
QHDU +D]HO YHQWUDO SHODJH ZLWK KDLUV 'HHS 1HXWUDO
*UD\EDVDOO\ZLWKVXSHU¿FLDO±PPZDVKRIOLJKW
FRORUHGIXU)LJ*XODUUHJLRQKDYLQJQRGLVWLQFWLYH
or contrasting markings. Fur of fore and hind limbs
colored like dorsum. Top of feet covered with Brown
hairs to base of metacarpals then replaced by longer,
VHOIFRORUHGKDLUV
Mystacial vibrissae present with longest extend
LQJWREDVHRISLQQDHZKHQODLGEDFNDORQJVLGHKHDG
superciliary vibrissae very sparse, about 12 mm long,
almost extending to dorsal edge of pinnae when laid
EDFN DORQJVLGH KHDG JHQDORQH YLEULVVDH SUHVHQW
interramal short. Ears sparsely covered with, brown
ish hairs, not contrasting conspicuously with color of
head. Opening to auditory canals protected by cteni
IRUPKDLUVSHVEURDGDOOGLJLWVZLWKXQJDOWXIWVRIVWLII
FWHQLIRUPEULVWOHVDQGVWURQJFODZVIRUHIRRWZLWKVKRUW
pollex bearing a broad open nail and digits II–V with
long, basally closed claws. Tail strong, darker above
WKDQEHORZLQPRVWVSHFLPHQVVSDUVHO\FRYHUHGZLWK
dark hairs, except for terminal 4–7 mm that are covered
with whitish hairs. Mammae not seen.
*ODQVSHQLVF\OLQGULFDOZLWKZHDNO\GHYHORSHG
EDVDOWURXJKDQGURXQGHGWLSXUHWKUDORSHQLQJYHQWUDO
SRVLWLRQHGDWWLSRISHQLVFRDUVHVSLQHVFRYHULQJJODQV
2VEDFXOXPPPORQJ/HVVDDQG&RRN
/F
6NXOOVPDOO)LJUREXVW ÀDWWHQHGLQGRUVDO
SUR¿OHLQWHURUELWDOUHJLRQZLWKURXQGHGPDUJLQV]\JR
matic arches parallel and not outwardly bowed, strongly
built, widest towards front of skull, with rounded
SUR¿OHLQGRUVDOYLHZPRUHHYLGHQWLQPDOHVGXHWR
VH[XDOGLPRUSKLVPRUDJHRUERWKSRVWRUELWDOSURFHVVHV
undeveloped and postorbital region posteriorly diver
JHQWDXGLWRU\EXOODHVOLJKWO\PRUHWKDQRIVNXOO
length, with auditory tubes long and salient (Fig. 9B,
E). Nasal bones broad, short. Interpremaxilary fora
men very small, inconspicuous. Frontals widest just
behind interorbital region, constricting just posterior to
HQGRISURFHVVHVRI]\JRPDWLFDUFKODPEGRLGDOULGJH
QRW ZHOOGHYHORSHG DOZD\V SUHVHQW HYHQ LQ \RXQJ
individuals. Supraoccipital crest strongly developed
in adults. Exoccipital portion of the ectotympanic re
duced. Mesopterygoid fossae with dorsal margin “V”
shaped, reaching to level between M2s. Posteropalatal
pits present, but not enlarged, palate grooved with deep
VXOFL $OLVSKHQRLG SUHVSKHQRLG YDULDEOH ÀDW DQG
broad or thin. Very narrow sphenopalatine vacuities.
,QFLVRUVVWURQJRUDQJHXSSHULQFLVRUVRUWKRGRQW0D[
illary tooth rows slightly divergent posteriad. Mandible
with coronoid process falciform, but not strongly
DQJOHGEDFNZDUGVFRQG\ORLGSURFHVVVWURQJEHDULQJ
DQDUWLFXODWLRQÀDQJHQRWZHOOGHYHORSHG
Karyology.—No karyotypes were prepared for
specimens of C. yatesi, all specimens were dead in
the traps.
Distribution and habitat.²(O WXTXLWR GH<DWHV
is known only from the type locality in the lowlands
of eastern Bolivia. The type locality is located in the
VRXWKHUQVHPLGHFLGRXV&KLTXLWDQRIRUHVWGLVWULFWRIWKH
Cerrado Biogeographic Province of eastern Bolivia
(Navarro and Maldonado 2002). Individuals were
collected from habitats with vegetation typical of the
&HUUDGR WKDW LQWHUGLJLWDWHV WKH /RZODQG &KLTXLWDQR
forest in this general area (see map in Cuéllar and Noss
7KLV &HUUDGR DOVR NQRZQ DV$ED\R\ LV
FKDUDFWHUL]HGE\WKHIROORZLQJSODQWVHymenaea stigonocarpa (Fabaceae), Luehea candicans (Tiliaceae),
%UHGHPH\HUD ÀRULEXQGD (Polygalaceae), Terminalia
argentea and T. fagifolia (Combretaceae), and Anacardium humile$QDFDUGLDFHDHRWKHUSODQWVSHFLHV
present in the area include Tabebuia selachidentata
1G$
$$$$$$$$$)''-%()!-,$&-&"4%=$8<%"<8$)*$6">-%$6"';$<!(."4%(69
(Bignoniaceae), Mimosa josephina (Fabaceae), and
Centratherum cardenasii$VWHUDFHDH
Etymology.—We name this species in honor of
RXUODWHIULHQGDQGJUHDWPHQWRU'U7HUU\/<DWHV
former curator of the Mammal Division of the Museum
of Southwestern Biology. His enthusiasm for system
DWLFVQDWXUDOKLVWRU\PXVHXPVDQG¿HOGPDPPDORJ\
ZDVDQGVWLOOLVDJUHDWVRXUFHRILQVSLUDWLRQWKDWZH
KRSHWRLQVWLOOLQRXUVWXGHQWV$Q\RQHZKRVKDUHGWLPH
LQWKH¿HOGZLWK7HUU\ZLOOUHFDOOKLV³UHPHPEHUWKH
motto” (“come back alive” and “save the specimens”)
H[SUHVVHGHTXDOO\DVDGHPDQGDVZHOODVDIDUHZHOOELG
(His inimitable Kentucky drawl permeates this passage
in our mind’s ear). The species name is a patronym in
the genitive singular.
Comparisons.—Only C. nattereri has been col
lected near the type locality of C. yatesi. Ctenomys
yatesi is much smaller (about 60–70% smaller) than
C. nattereriLQDOOPHDVXUHPHQWVWDNHQKRZHYHUWKH
VL]HRIWKHW\PSDQLFEXOODHUHODWLYHWRVNXOOVL]HLV
larger in C. yatesi than in C. nattereri. In addition,
the dorsal coloration in C. yatesi is lighter with weak
differentiation between dorsum and venter with no
collar versus a darker, Chocolate Brown above with a
FRQWUDVWLQJ<HOORZYHQWHUDQGDYHU\GLVWLQFW<HOORZ
collar in C. nattereri. Ctenomys yatesi differs from C.
steinbachiLQEHLQJVPDOOHUZLWKWKHGRUVDOIXU+D]HOLQ
color (rather than Fuscous Black), and ventrally Light
+D]HOUDWKHUWKDQ*UL]]OHG:KLWLVKDQG)XVFRXVLQ
addition, C. yatesi has a relatively longer tail (compared
to head and body length), and narrower interorbital
breadth (Table 1).
Ctenomys yatesi differs from C. conoveri in be
ing much smaller in all measurements. In addition,
C. yatesiKDVDWLG\SHODJHWKDWLV+D]HOLQFRORUDQG
differs dramatically from the shaggy Brownish pelage
of C. conoveri.
Ctenomys yatesi can be differentiated from C.
boliviensis, especially the populations to the south of
6DQWD&UX]HJ/DV/RPLWDVLQFRPSOHWHO\ODFNLQJD
EODFNFDSDQGVWULSHWKDWUXQVPLGGRUVDOO\WRWKHUXPS
in C. boliviensis. In C. yatesi the ventral coloration is
less Ochraceous than in C. boliviensis and the skull
is 50 to 65% smaller, with a very different shape of
the tympanic bulla and rostrum. Ctenomys from the
highlands or dry valleys of Bolivia occur in habitats
very distinct from that of type locality of C. yatesi and
DOODUHPXFKODUJHULQERG\VL]H
2WKHU VSHFLHV RI VPDOOERGLHG Ctenomys in
clude C. bergi and C. pundti IURP$UJHQWLQD 2OURJ
and Lucero 1981) and C. dorsalis from Paraguay
7KRPDV 5HGIRUG DQG (LVHQEHUJ 7KH
ranges of C. bergi and C. pundtiLQ&yUGREDDQG6DQ
/XLVSURYLQFHVRIQRUWKFHQWUDO$UJHQWLQDDUHIDUIURP
that of C. yatesi and the biomes are distinctly different.
6HTXHQFHGLYHUJHQFHLVEHWZHHQC. pundti and
C. yatesi (no comparable data exist for C. bergi, but a
VKRUWIUDJPHQWRIESVKRZHGGLYHUJHQFHZLWK
DQKRPRORJRXVIUDJPHQWLQ<DWHV¶WXFRWXFR7KHVH
are values higher than those suggested by Lessa and
&RRNIRUJHQHWLFGLYHUJHQFHEHWZHHQVSHFLHV
SDLUVRIWXFRWXFRVCtenomys yatesi differs from C.
dorsalis D VPDOOERGLHG VSHFLHV IURP WKH &KDFR RI
Paraguay, in the very distinct coloration pattern of
their body pelages: C. dorsalis has a distinct collar of
light hairs behind the cheeks which extends to the ears
RQHDFKVLGHRIWKHKHDGLQDGGLWLRQWKLVVSHFLHVKDV
a stripe of black hairs on the head, which runs down
the back of the body and most of the ventral side of
WKHERG\LVFRYHUHGZLWKVHOIFRORUHGKDLUVRUDUHSDOH
buffy (Thomas 1900). Ctenomys yatesi is, in contrast,
rather uniform in coloration, lacks the stripe of dark
hairs on the back and has Buffy or in some instances
ZKLWLVKWLSSHGKDLUVLQWKHYHQWHUZKLFKDUHRWKHUZLVH
VODWHFRORUHGDWWKHLUEDVH
*HQHWLFDOO\C. yatesi differs from all known spe
cies in the lowlands of Bolivia from a low of 5.4% to
a high of 10.2% (Table 1).
Remarks.²$QGHUVRQHWDOXVHGWKHQDPH
C. minutus that Nehring coined in 1887 to refer to these
specimens as “a reasonable working taxonomic hy
SRWKHVLVDOWKRXJKTXLWHWHQWDWLYH´$QGHUVRQDQGFRO
ODERUDWRUVMXVWL¿HGXVLQJWKHQDPH³minutus” because
&DEUHUDKDGV\QRQ\PL]HGC. bicolor (Miranda
5LEHLURDVPDOOERGLHGIRUPIURPWKH%UD]LOLDQ
VWDWHRI5RQG{QLD%LGDXDQG$YLOD3LUHVZLWK
C. minutus. Ctenomys minutusLVDOVRDVPDOOERGLHG
VSHFLHVRULJLQDOO\GHVFULEHGIURPVRXWKHDVWHUQ%UD]LO
5LR*UDQGHGR6XO&DPSRV(RI0RQGR1RYR&RP
parisons of C. bicolor with other species of Ctenomys
IURP%UD]LOE\6WROW]HWDOXVLQJPRUSKRORJLFDO
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
FKDUDFWHUVFKURPRVRPHVDQGJHQHWLFVFRQ¿UPWKHYD
lidity of C. bicolor as a distinct species that has closer
DI¿QLWLHVWRC. boliviensis than to C. minutus.
Ctenomys yatesi GLIIHUVLQVL]HDQGLQFRORUDWLRQ
from C. minutus, and these two species also have
UDQJHV WKDW DUH ZHOOVHSDUDWHG LQ JHRJUDSKLF VSDFH
Morphologically, C. yatesi and C. minutus differ in
many characters with C. yatesi having a shorter ros
WUXPODUJHUDQGPRUHLQÀDWHGDXGLWRU\EXOODHVKRUWHU
and broader nasals, and smaller paroccipital apophyses
RQWKHW\PSDQLFEXOODH)LQDOO\EXWVLJQL¿FDQWO\ZH
measured genetic divergence of cytE VHTXHQFHV EH
tween these species at 5.8%.
Ctenomys andersoni, new species
$QGHUVRQ¶V&XMXFKL(OFXMXFKLGH$QGHUVRQ
)LJVPDS7DEOH
Ctenomys “ita”: Lessa and Cook (1998:89)
Ctenomys “ita”: Cook and Lessa (1998:1,524)
C. sp. “Ita”: Castillo et al. (2005:172)
Ctenomys VS,7$3DUDGDHWDO
Holotype.²06%0XVHXPRI6RXWKZHVW
ern Biology), adult male. Skin, skull and axial skeleton
FROOHFWHG-XO\SUHSDUHGE\6\GQH\$QGHUVRQ
¿HOGQXPEHU6$$WWKHWLPHRIFROOHFWLRQWKH
OLYHUNLGQH\DQGKHDUWWLVVXHVZHUHIUR]HQLQOLTXLG
QLWURJHQDQGVXEVHTXHQWO\VWRUHGDWƒ&LQWKH'LYL
VLRQRI*HQRPLF5HVRXUFHV0XVHXPRI6RXWKZHVWHUQ
Biology, NK21221.
Type locality.—Bolivia, Department of Santa
&UX] &HUUR ,WDKXDWLFXD °
6 °
: P
elevation (see map, Fig. 8).
Measurements of holotype.²727/7$,/
+)/($5:7J&%5&,/
3/502/'6%3$%=<%
,2%/$%6+7
Specimens examined.²7ZHQW\HLJKWLQGLYLGXDO
specimens, here designated as paratypes, all from the
W\SHORFDOLW\$01+
VHYHQ
06% VHYHQ01.1
1/
625). Pooled measurements of skulls and external
GLPHQVLRQVRIERWKVH[HVDUHVXPPDUL]HGLQ7DEOH
VHH7DEOH6 LQ RQOLQH VXSSOHPHQWDO GRFXPHQWV IRU
FRPSOHWHGDWDVHW
Diagnosis.²$ PHGLXPVL]HG PHPEHU RI WKH
“boliviensis” JURXS RI WXFRWXFRV ZLWK EURZQ GRUVDO
coloration, an almost indistinct Olive Brown stripe, and
QRFDSRIGDUNKDLUVRQKHDGYHQWHUPXFKOLJKWHUDQG
ZHOOGLIIHUHQWLDWHGIURPGRUVDOFRORUDWLRQ1RGDUNFRO
ODULQJXODUUHJLRQVPDOOSDWFKHVRIVHOIFRORUHGKDLURQ
pectoral, inguinal, jowl and/or axillary regions. Skull
with a short and narrow rostrum, very short nasals,
SURRGRQWLQFLVRUV]\JRPDWLFSURFHVVPRVWO\VWUDLJKW
and a karyotype with 2N=46 and FN=50 of mostly
acrocentric autosomes.
Description (Figs. 11, 12).—Dorsal pelage
dense, buffy brown to Mummy Brown darkening to
an indistinct light Olive Brown stripe starting at the
KHDGSURFHHGLQJSRVWHULDGIXUUHODWLYHO\FRDUVHDERXW
WRPPORQJRYHUEDFNDQGUXPSKDLUVDQGIXUDW
EDVHDGHHS0RXVH*UD\RU1HXWUDO*UD\H[FHSWIRUWKH
distal most part of each hair that is a distinctly lighter
VKDGHRI%XII\%URZQ*XODUUHJLRQOLJKWHULQFRORU
and with a wash of Pale Olive Buff under forelegs.
Color of ventral fur lighter than dorsal fur, Pale Olive
%XIIZLWKKDLUVD1HXWUDO*UD\RU'HHS1HXWUDO*UD\
EDVDOO\ZLWKVXSHU¿FLDO±PPZDVKRI3DOH2OLYH
Buff or Buffy Brown. Inguinal area light pale Olive
Buff in color, lighter than ventral fur in general. Top of
feet covered with hairs to base of metacarpals replaced
with cteniform hairs over each toenail and on sides of
toes. Some mystacial vibrissae extending past pinnae
ZKHQODLGEDFNDORQJVLGHKHDGVXSHUFLOLDU\YLEULVVDH
VSDUVH H[WHQGLQJ WR WKH EDVHV RI SLQQDH JHQDORQH
present, interramal present.
Ears sparsely covered with short, brownish hair,
contrasting slightly with color of head. Opening to
auditory canals protected by conspicuous cteniform
bristles, most individuals with white patches of fur
EHORZ DQG SRVWHULRU WR SLQQDH EXQGOHV RI VWLII KDLU
posterior to upper and lower incisors. Pes broad, all
digits with ungal tufts of stiff “cteniform” bristles and
VWURQJFODZVFWHQLIRUPEULVWOHVRQPLGGOHFODZDERXW
KDOIDVORQJDVQDLO)RUHIRRWZLWKDZHOOGHYHORSHG
pollex bearing a short nail open basally in contrast to
the other four digits that bear basally closed claws. Tail
11$
$$$$$$$$$)''-%()!-,$&-&"4%=$8<%"<8$)*$6">-%$6"';$<!(."4%(69
)LJXUH6NXOODQGPDQGLEOHRIKRORW\SH06%RICtenomys andersoniQVS$GRUVDOYLHZRIVNXOO%
ventral view of skull, C) dorsal view of mandible, D) ventral view of mandible, E) en face view of skull showing dark
RUDQJHFRORURIXSSHULQFLVRUV)OHIWODWHUDOYLHZRIVNXOO*OHIWODWHUDOYLHZRIPDQGLEOH
)LJXUH0XVHXPVWXG\VNLQRIKRORW\SH06%RICtenomys andersoniQVS$GRUVDOYLHZ%YHQWUDO
view, C) left lateral view.
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
1B
)LJXUH,PDJHRIVWDQGDUGJLHPVDVWDLQHGNDU\RW\SHRICtenomys andersoniQVS06%FRQVLVWLQJRI
19 pairs of acrocentric autosomes, two pairs of metacentric autosomes, and one pair of submetacentric autosomes.
7KHVH[FKURPRVRPHVLQER[DSSHDUDVDODUJHVXEPHWDFHQWULF;DQGDQDFURFHQWULF<7KH1 )1 VKRUWVWURQJWHUHWLIRUPLQFURVVVHFWLRQGDUNHUDERYH
WKDQEHORZLQPRVWVSHFLPHQVVSDUVHO\FRYHUHGZLWK
dark hairs, except for terminal 4 mm, which are covered
with whitish hairs.
6NXOO UREXVW )LJ LQWHURUELWDO UHJLRQ ZLWK
PRVWO\VTXDUHGPDUJLQV]\JRPDWLFDUFKHVEURDGDX
GLWRU\EXOODHLQÀDWHGS\ULIRUPDXGLWRU\WXEHVORQJ
VDOLHQW 1DVDO ERQHV EURDG EURDGHVW WKUHHIRXUWKV
DQWHULDGZLWKÀDWGRUVDOHGJHVQDVDOVKRUWSUHPD[LO
lary visible when viewed from above. Frontals widest
just at the middle of interorbital region, constricting
at the same level as the attachment of the rear part of
WKH]\JRPDWLFDUFKSURFHVVHVSDULHWDOVLQROGHULQGL
YLGXDOVQDUURZHVWDWDWWDFKPHQWRIVTXDPRVDOURRWRI
]\JRPDWLFDUFK6TXDPRVDOHYLGHQWO\H[SDQGLQJZLWK
DJHLQ\RXQJLQGLYLGXDOVSDULHWDODQGUHDURIIURQWDOV
are approximately the same width. Interparietal com
pletely fused in older individuals. Temporal foramen
DWMXQFWLRQRIVTXDPRVDODQGSDULHWDODOZD\VSUHVHQW
Lambdoidal ridge always present. Most individuals
ZLWKIHQHVWUDEHWZHHQIURQWDOVDQGSDULHWDOV*DUGQHU
DQG$QGHUVRQ 6XSUDRFFLSLWDO FUHVW VWURQJO\
GHYHORSHG LQ DGXOWV =\JRPDWLF DUFK VWURQJO\ EXLOW
MXJDOVZLWKVWURQJGRUVRYHQWUDOSURFHVVHVLQFLVLYHIR
ramina recessed in a common fossa and well separated
E\DVWURQJO\GHYHORSHGERQ\VHSWXPPRVWLQGLYLGXDOV
ZLWKDVPDOOLQWHUSUHPD[LODU\IRUDPHQSDODWDOEULGJH
with two major palatine foramina at level of M1. Me
sopterygoid fossae, an inverted “V” shape, reaching
anteriorly to middle of M2s. Posteropalatal pits pres
HQWEXWQRWHQODUJHG$OLVSKHQRLGSUHVSKHQRLGEULGJH
1C$
$$$$$$$$$)''-%()!-,$&-&"4%=$8<%"<8$)*$6">-%$6"';$<!(."4%(69
narrow and with evident lateral processes. Bony roof
ZLWKVSKHQRSDODWLQHYDFXLWLHVEXFFLQDWRUPDVWLFDWRU\
foramen enlarged and undivided in many individuals,
although in the holotype these foramina are separated.
Upper incisors large, robust, Orange in color, proodont.
Maxillary tooth rows slightly divergent posteriad.
Mandible robust, with coronoid process falci
IRUPDQJOHGVWURQJO\EDFNZDUGVFRQG\ORLGSURFHVV
VWURQJEHDULQJDZHOOGHYHORSHGDUWLFXODWLRQÀDQJH
Karyology.²1 )1 )LJ ZLWK pairs of acrocentric autosomes, two pairs of metacentric
autosomes, and one pair of submetacentric autosomes.
The sex chromosomes appear as a large submetacentric
;DQGDODUJHDFURFHQWULF<$VHFRQGDU\FRQVWULFWLRQ
is evident on the largest biarmed pair of chromosomes.
Karyotype slides were prepared in the field from
QLQH VSHFLPHQV LQFOXGLQJ WKUHH PDOHV 06%
$01.DQGVL[IHPDOHV$0
06%06%01.101.1DQG
NK21259).
Distribution and habitat.²$QGHUVRQ¶V&XMXFKLLV
known only from the type locality at Cerro Itahuaticua.
Here the vegetation was a mixture of deciduous thorny
trees (Fabaceae) and several species of cacti, primarily
Opuntia spp. Limestone outcrops were common on
KLOOVLGHV7KLVORFDOLW\LVLQWKHHFRORJLFDO]RQHNQRZQ
DV ERVTXHV VHFRV LQWHUDQGLQR ,ELVFK HW DO RU
$QGHDQGU\YDOOH\V8Q]XHWD/RSH]
Etymology.—This species of Ctenomys is named
LQKRQRURI'U6\GQH\$QGHUVRQ&XUDWRU(PHULWXVRI
WKH'HSDUWPHQWRI0DPPDORJ\$PHULFDQ0XVHXPRI
1DWXUDO+LVWRU\6WDUWLQJLQ'U$QGHUVRQOHDG
D WHDP RI LQYHVWLJDWRUV FKLHÀ\ IURP$01+ 06%
CBF, and MNKM, on collecting expeditions throughout
Bolivia, including trips that secured these specimens.
$ JUHDW PHQWRU DQG D GHDU FROOHDJXH ZH UHPHPEHU
him examining the day’s catch and then preparing
specimens for hours on end. We are pleased to name
WKLV%ROLYLDQHQGHPLFWRKRQRUKLVOHJDF\WKHUHIRUH
the name is a patronym in the genitive singular and in
apposition to the generic name.
Comparisons.—Two specimens show a weak
collar of pale fur extending ventrad from the pinnae,
but not to the extent seen in C. boliviensis, which has
a very well developed gular collar of light colored fur.
Ctenomys andersoni n. sp. differs from C. steinbachi, a species known from lowland Bolivia near Santa
&UX]E\KDYLQJQRJXODUFROODURIOLJKWFRORUHGIXUE\
having fur that is comparably lighter in color and by
having a diploid and fundamental chromosome number
of 2N=46, FN=50 versus 2N=10, FN=16 in C. steinbachi VHH$QGHUVRQHWDO,QDGGLWLRQWKHNDU\R
type of C. andersoni has mostly acrocentric somatic
FKURPRVRPHVZLWKRQO\WKUHHELDUPHGSDLUVZKLOHWKH
chromosomes of C. steinbachi are all biarmed.
Ctenomys andersoni is distinct from C. frater,
which occurs in the eastern foothills of Bolivia, by
having light Mummy Brown fur instead of Dark Brown
RU1HXWUDO*UD\IXUDVLQC. frater and by chromosome
numbers (2N=48 and FN=90 in C. frater) see Cook et
al. (1990).
Ctenomys andersoni is much smaller than C.
conoveri, has much shorter fur, and has a diploid
chromosome number of 2N=46 versus 2N=48 in C.
conoveri VHH$QGHUVRQHWDO
The only other species occurring in the geograph
ic proximity of C. andersoni is C. lewisiVSHFLPHQVRI
C. andersoni were collected in much different habitat
and much lower altitude than C. lewisi, which is known
RQO\IURPDQDUHDQRUWKZHVWRIWKHFLW\RI7DULMDDWHO
HYDWLRQVDERYHP,QDGGLWLRQWKHGRUVDOERG\IXU
of C. andersoni is brown in color, relative to C. lewisi,
which is dark (Fuscous Black) dorsally. Chromosome
numbers also differ between these species (2N=56,
FN=74 in C. lewisi; Cook et al. 1990).
Finally, C. andersoni is genetically distinct from
all other species in and around Bolivia (see Table 2)
with genetic distances ranging from 5.0% in compari
son with C. erikacuellarae to approximately 10% with
both C. lewisi and C. frater. Other species of Ctenomys
from Bolivia need not to be compared directly with C.
andersoni as their ranges include high altitude altiplano
and puna habitat (C. opimus and C. leucodon).
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
Ctenomys lessai, new species
/HVVD¶V7XFR7XFR(OWXFRWXFRGH/HVVD
Figs. 8 (map), 14, 15, 16, Table 1
Ctenomys “llathu”: Lessa and Cook (1989:89)
Ctenomys “llath”: Cook and Lessa (1998:1,524)
Ctenomys “llathu”: Lessa and Cook (1998:89)
Ctenomys “Llathu”: Castillo et al. (2005:172)
Ctenomys sp. Llathu: Parada et al. (2011:682)
C.VS//$7+8)UHLWDVHWDO
Holotype.—MSB67111 (Museum of South
western Biology), adult female. Skin, skull, and axial
VNHOHWRQFROOHFWHG0D\SUHSDUHGE\)RUHVW
:'DYLV¿HOGQXPEHU):'7KHOLYHUNLGQH\
DQGKHDUWWLVVXHZHUHIUR]HQDQGDUFKLYHGLQWKH'LYL
VLRQRI*HQRPLF5HVRXUFHVRIWKH06%ZLWKQXPEHU
NK22870.
Type locality.—Bolivia, Department of Co
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6
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:PHOHYDWLRQ
Measurements of holotype.²727/7$,/
+)/ ($5 :7 J &%5 &,/3/502/'6%3$%
=<%,2%/$%6+7
Specimens examined.—Six specimens desig
nated as paratypes from the type locality: Two MSB
IRXU$01+
264560). Two, MNKN (two specimens, no museum
FDWDORJQXPEHUVDYDLODEOH¿HOGQXPEHUVDQG06%
'*5 FDWDORJ QXPEHUV 1. 1. QRW
paratypes. Pooled measurements of skulls and external
GLPHQVLRQVRIERWKVH[HVDUHVXPPDUL]HGLQ7DEOH
VHH7DEOH6 LQ RQOLQH VXSSOHPHQWDO GRFXPHQWV IRU
WKHFRPSOHWHGDWDVHW
Diagnosis.²$ PHPEHU RI WKH ³frater” group
of CtenomysFKDUDFWHUL]HGE\DGRUVDO2OLYH%URZQ
coloration with a dark cap of Clove Brown hair cover
ing most of the head. Pectoral region colored same
DVGRUVXPEXWYHQWHUFRYHUHGZLWKVHOIFRORUHGKDLUV
No collar in gular region. Skull delicate with mostly
VWUDLJKW]\JRPDWLFDUFKHVDQGDÀDWVNXOOSUR¿OHLQFL
sors proodont, with pale yellow enamel and a karyotype
with 2N=46 and FN=64.
1D
Description.²3HODJH GHQVH ¿QH VRIW DERXW
5–20 mm long over back and rump. Color of dorsal
pelage Olive Brown to Buffy Brown, ventral pelage
Cinnamon Buff, some individuals with ventral fur Olive
Buff. Dorsally, darkest fur a Clove Brown in central
diffuse dorsal stripe, more prominent anteriorly on
head and fading posteriad (Fig. 14). Small light area
of Cinnamon Buff fur just posterior to and below pin
nae, evident in most individuals. Collar not evident in
VSHFLPHQVH[DPLQHG*XDUGKDLUVRIYDU\LQJOHQJWKV
approximately twice length of undercoat. Undercoat
IXU ELFRORUHG ORZHU ô RI KDLUV GHHS 1HXWUDO *UD\
distal parts with color as described above. Juveniles
ZLWKFRQVLVWHQWGHHS1HXWUDO*UD\FRORUQRELFRORUHG
PDUNLQJV0\VWDFLDOYLEULVVDHRIYDULRXVVL]HVIURP
anterior to posteriad, initial vibrissae shorter, medial
vibrissae longest in group, extending past base of pin
QDHZKHQODLGSRVWHULDGGLVWDOYLEULVVDHVKRUWHU*HQDO
vibrissae present, interramal present. Ears sparsely
furred, same color as head, inconspicuous cteniform
hairs protecting ear canal. Bundles of stiff hair on
lips posterior to both upper and lower incisors. Rear
legs with broad pes, all digits with ungal tufts of stiff
FWHQLIRUPEULVWOHVIRUHIRRWZLWKZHOOGHYHORSHGGLJLWV
bearing basally closed claws. Tail short, covered with
dark Olive Brown hair, teretiform, lighter fur ventrally,
end of tail lighter in color.
Skull relatively delicate (Fig. 15), interorbital
UHJLRQZLWKURXQGHGPDUJLQV]\JRPDWLFDUFKHVZLGH
extending laterally almost to level of external audi
WRU\ PHDWXV $XGLWRU\ EXOODH S\ULIRUP QDVDO ERQHV
broad tapering to a “V” shape posteriad. Frontal bones
broad, constricting to narrowest point just posterior
WR QDVDOIURQWDO VXWXUH 3DULHWDOV QDUURZHVW DW SRLQW
RI DWWDFKPHQW RI VTXDPRVDO URRW RI ]\JRPDWLF DUFK
Interparietal not visible. Temporal foramen at junc
WLRQRIVTXDPRVDODQGSDULHWDODOZD\VSUHVHQWRYRLG
Lambdoidal ridge present, more pronounced in older
individuals. Zygomatic arch strong, with massive pro
cesses extending both dorsally and ventrally. Incisive
foramina recessed in a common fossa and incompletely
separated by a bony septum at junction of premaxillary
and maxillary bones. Small interpremaxillary foramen
FRQVSLFXRXVVPDOOSDODWLQHIRUDPLQDVRPHWLPHVRII
VHWVRPHWLPHVSDUDOOHODWOHYHORI30DQG0DQWHULRU
edge of mesopterygoid fossae with strongly inverted
9VKDSHGERUGHUUHDFKLQJWROHYHORIDQWHULRUSDUWRI
12$
$$$$$$$$$)''-%()!-,$&-&"4%=$8<%"<8$)*$6">-%$6"';$<!(."4%(69
)LJXUH0XVHXPVWXG\VNLQRIKRORW\SH06%RICtenomys lessaiQVS$GRUVDOYLHZ%YHQWUDOYLHZ
C) left lateral view.
Figure 15. Views of the skull and mandible of the holotype (MSB67111) of Ctenomys lessaiQVS$GRUVDOYLHZ
of skull, B) ventral view of skull, C) dorsal view of mandible, D) ventral view of mandible, E) en face view of skull
VKRZLQJ\HOORZLVKFRORURIXSSHULQFLVRUV)OHIWODWHUDOYLHZRIVNXOO*OHIWODWHUDOYLHZRIPDQGLEOH
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
1E
)LJXUH,PDJHRIVWDQGDUGJLHPVDVWDLQHGNDU\RW\SHRICtenomys lessai n. sp. consisting of one male
06%DQGRQHIHPDOH$0>1 )1 @LQFOXGLQJVHYHQSDLUVRIVXEWHORFHQWULFWZR
SDLUVVXEPHWDFHQWULFSDLUVDFURFHQWULFRQHSDLURIYHU\VPDOOGLVWLQFWLYHVXEWHORFHQWULFDQGRQHSDLU
RIDFURFHQWULFVH[FKURPRVRPHV$ IHPDOH% PDOH
M2. Suture of basioccipital and basisphenoid located
at site of attachment of auditory bullae. Buccinator
and masticatory foramina joined and enlarged even
in young individuals. Upper incisors with enamel a
OLJKWSDOH<HOORZ2FKHUWR,YRU\<HOORZFRORU(QDPHO
LQLWLDOO\ ZKLWH LQ XQH[SRVHG URRWV FKDQJLQJ WR<HO
low Ocher with mottled yellowing slightly on frontal
surfaces of exposed parts of incisors. Sometimes ap
pearing mottled in frontal view with white showing
WKURXJK(QDPHORIORZHULQFLVRUV<HOORZ2FKHULQ
color. Mandible relatively delicate. Flange of mastoid
process not visible.
Distribution and habitat.—Known only from the
type locality with specimens collected at elevations
ranging from 2,500 to 2,750 m. The area was an open
grassland habitat near a running stream with remnant
stands of trees (Polylepis sp.).
Karyology.—Two animals from the type locality
were karyotyped (one male, MSB67111 and one female
$0ZLWK1 )1 )LJ.DU\RW\SH
consisting of chromosomes that include: seven pairs
RIVXEWHORFHQWULFWZRSDLUVVXEPHWDFHQWULFSDLUV
DFURFHQWULFRQHSDLURIYHU\VPDOOGLVWLQFWLYHVXEWHOR
centric, and one pair of acrocentric sex chromosomes.
Comparisons.—Ctenomys lessai differs from C.
erikacuellaraeWKHVSHFLHVRIWXFRWXFRZLWKWKHFORV
est geographic range to the southeast (see map, Fig. 8),
LQKDYLQJSHODJHRIDGDUNHUDQGPRUHXQLIRUPFRORU
upper incisors smaller, more procumbent, and much
lighter in color than in C. erikacuellarae$OVRLQC.
lessai, the lateral basisphenoid foramina are conspicu
Etymology.—This species of Ctenomys is named
LQKRQRURI'U(QULTXH3/HVVDDIULHQGJLIWHGPHQ
tor, and talented gaucho who has emerged as a leader
LQ/DWLQ$PHULFDQPDPPDORJ\HYROXWLRQDQGLQWKH
VWXG\RIWKHELRORJ\RIWXFRWXFRV7KHVSHFLHVQDPH
is thus a patronym in the genitive singular.
10$
$$$$$$$$$)''-%()!-,$&-&"4%=$8<%"<8$)*$6">-%$6"';$<!(."4%(69
ously enlarged but are either very small or not evident
in C. lessai or in C. erikacuellaraeDQGWKHH[WHUQDO
auditory meatus is directed more anteriorly compared
to C. erikacuellarae. Species of Ctenomys occurring
farther east and at lower elevations, including C. an-
dersoni, C. boliviensis, and C. steinbachi, are all larger
LQERG\VL]HZLWKGLIIHUHQWSHODJHFRORUDQGDOOKDYH
different karyotypes. Finally, genetic distances based
on cytb range from 6.8% with C. conoveri to 11.6%
with C. leucodon (Table 1).
5(%'<%%()!
Recognition of Ctenomys goodfellowi as a junior
synonym of C. boliviensis.—Originally described by
7KRPDVIURP(VSHUDQ]DQHDU&RQFHSFLyQLQ
eastern Bolivia, the status of C. goodfellowi has histori
FDOO\ÀXFWXDWHGIURPEHLQJFRQVLGHUHGDVXEVSHFLHVRI
C. boliviensis &DEUHUD $QGHUVRQ HW DO *DUGQHU WR EHLQJ UHFRJQL]HG DV D IXOO VSHFLHV
&RRN DQG <DWHV $QGHUVRQ 7KRPDV
LGHQWL¿HG VXEWOH PRUSKRORJLFDO GLIIHUHQFHV
that served to separate the single individual that he
described as C. goodfellowi from one of C. boliviensis
(which was then designated as a lectotype of C. boliviensis$QGHUVRQHWDOUHSRUWHGD1 IRU
individuals assigned to C. goodfellowi and compared
both morphological and chromosomal characters for
individuals assigned to both species concluding that
these may represent population level variation. In his
SK\ORJHQ\EDVHGRQO\RQPRUSKRORJ\*DUGQHU
showed that the subspecies C. boliviensis boliviensis
Waterhouse 1848 and C. boliviensis goodfellowi (along
with C steinbachi) are derived from a common ancestor,
thus providing some support to hypothesis that these
populations of C. boliviensis probably represented a
single species.
Based on electromorphic variation at 21 presump
WLYHORFL&RRNDQG<DWHVFRPSDUHGLQGLYLGX
als from four chromosomally polytypic populations
that were assumed to all represent C. boliviensis LQ
WKLVDQDO\VLVWKH\IRXQGDPLQRUDOOHOLFIUHTXHQF\
variations among most samples of the species, with
H[FHSWLRQ RI DOOHOH 62'E WKDW ZDV ¿[HG LQ WKH C.
boliviensisSRSXODWLRQIURP/DV/RPLWDVEXWLPSRU
tantly, b) one population (C. boliviensis, SRSXODWLRQ
WKDWZDVFRPSRVHGRILQGLYLGXDOVZLWKD1 WKDW
ZDVTXLWHGLIIHUHQWDQGSUHVHQWHGXQLTXH¿[HGDOOHOHV
when compared to the remaining populations in their
samples of C. boliviensis. Our phylogenetic analyses
)LJV DQG VKRZ WKDW WKHVH SRSXODWLRQV VSHFL¿
cally, (C. boliviensisSRSXODWLRQRI&RRNDQG<DWHV
(1994) belong to C. nattereri$WWKHWLPH&RRNDQG
<DWHVVXJJHVWHGWKDWC. goodfellowi deserved
VSHFLHVOHYHO UHFRJQLWLRQ EDVLQJ WKLV UHFRPPHQGD
tion on their study of both chromosomes (2N=46) and
DOOR]\PH GLIIHUHQFHV DOOHOH IUHTXHQF\ GLIIHUHQFHV
$QGHUVRQ/HVVDDQG&RRN&RRNDQG
Lessa (1998), Parada et al. (2011), Freitas et al. (2012)
KDYHVLQFHIROORZHGWKHFRQFOXVLRQRI&RRNDQG<DWHV
(1994).
The status of C. goodfellowi, however, is predi
cated on a clearer understanding of the nature of the
entire “boliviensis” group of Ctenomys, something that
is only now emerging. Relevant to this discussion is
the status of three species with geographic distributions
PRVWO\LQ%UD]LOVHH)LJWKDWLQFOXGHCtenomys
nattereri, C. rondoni, and C. bicolor and are species that
are poorly represented in natural history collections.
5HFHQWO\6WRO]HWDOXVHGJHRPHWULFPRUSKR
PHWULFDQDO\VHVDQGSDUWLDOVHTXHQFHVRIcytb to validate
the status of C. bicolor as distinct from C. nattereri.
2SSRUWXQHO\ WKHVH DXWKRUV VDPSOHG TXDVLWRSRW\SL
cal material assigned to the latter, which in turn also
provides support for the status of C. nattereri as a full
VSHFLHV:LWKWKHVHWZRVSHFLHVPRUHFOHDUO\GH¿QHG
the nature of C. goodfellowi can now be better under
stood as follows: 1) based on the fact that individuals
assigned to C. goodfellowi consistently are recovered
as paraphyletic with respect to C. boliviensis in our
phylogenies, 2) that their diploid chromosome number
appears to be part of a cline of variation (from 2N=42
to 2N=46) in populations assigned to C. boliviensis
(which includes the presence of at least one 2N=45 in
GLYLGXDODQGYHU\VPDOOYDOXHVRIJHQHWLFGLVWDQFHV
between populations assigned to C. goodfellowi and C.
boliviensisLQFRPSOHWHcytEVHTXHQFHGDWD
LQHOHFWURSKRUHWLFGDWD:HWKHUHIRUHV\QRQ\PL]HC.
goodfellowi under C. boliviensis. We acknowledge that
genetic and morphological variation among populations
assigned to C. goodfellowi and C. boliviensis exist,
?-45!"4$"6$-,@A!"#$%&"'("%$)*$!"#$%&'($*4)8$+),(.(-$
EXWVXJJHVWWKDWWKHVHGRQRWIXO¿OOWKHFULWHULDZHVHW
in the introduction of this report for the recognition of
species in the genus.
Our phylogenetic analyses, the most inclusive to
date for species in the genus Ctenomys, show that the
“boliviensis” group of Ctenomys (sensu Parada et al.
2011) is only weakly supported and is composed of
two groups of species: One that includes C. bicolor,
C. nattereri, C. boliviensis, and C. steinbachi and one
that includes three of the four species described in this
report: C. erikacuellarae, C. yatesi, and C. andersoni.
The importance of the Andes as a speciation engine in Ctenomys.—The complicated orogeny of the
$QGHVKDVFUHDWHGLQ%ROLYLDQXPHURXVULGJHVYDOOH\V
DQG FDQ\RQV VWUHWFKLQJ LQ D QRUWKVRXWK RULHQWDWLRQ
DORQJLWVHDVWHUQÀDQN3DUWRIWKH<XQJDVDQGWKH$Q
dean dry valleys in the Departments of Cochabamba,
6DQWD&UX]&KXTXLVDFDDQG7DULMDKDYHDJHRPRUSKLF
RULJLQFRWHPSRUDOZLWKDK\SRWKHVL]HGUDSLGXSOLIWRI
WKHFHQWUDO$QGHVEHWZHHQaDQGPLOOLRQ\HDUVDJR
0\D+RUWRQHWDO*DU]LRQHHWDO(KOHUV
DQG3RXOVHQ7KH$QGHVWRSRJUDSK\FRPELQHG
ZLWKVORSHDQGDVSHFWDUHUHVSRQVLEOHIRUKLJKO\XQLTXH
microclimatic and ecological conditions, which in some
FDVHVFKDQJHDFURVVYHU\VKRUWGLVWDQFHV$ODWLWXGLQDO
transect (ca. 18°6WKURXJKWKHFHQWUDOSODWHDXRIWKH$Q
des shows the extreme topographic relief in the central
$QGHDQEDFNWKUXVWEHOWVHH¿JXUHVLQ0F4XDUULHHWDO
DQG)LJ)LQRQOLQHVXSSOHPHQWDOGRFXPHQWV
The orogeny of the mountains and formation
of valleys in this belt have been implicated as drivers
RIELRGLYHUVLW\LQWKHFHQWUDO1HRWURSLFV*UDKDPHW
DO+XJKHVDQG(DVWZRRG7KHUHJLRQ¶V
FKDUDFWHUL]DWLRQDVDKRWVSRWRIELRGLYHUVLW\FDQEHDW
tributed partially to the combination of microclimatic
variability and changing environmental conditions
WKURXJK WKH 4XDWHUQDU\ 7KH PXOWLSOH <XQJDV DQG
$QGHDQGU\YDOOH\VDQGDVVRFLDWHGQRUWKVRXWKULGJHV
likely act as geographic barriers to movement of organ
isms. These barriers and microclimatic variability may
HVSHFLDOO\VWLPXODWHVSHFLHVGLYHUVL¿FDWLRQLQVXEWHUUD
nean rodents (Nevo 1979). Ongoing formation of new
ULGJHVIURPDFWLYHRURJHQLFSURFHVVHV0F4XDUULHHW
al. 2005) separate populations in the area, leading to a
GHFUHDVHLQHDVWZHVWJHQHÀRZZKLOHDOORZLQJQRUWK
1F
VRXWKGLVSHUVDOWRFRQWLQXHVHH)LJV))DQG)
in online supplemental documents).
The Octodontidae and the Ctenomyidae appear
to have originated from a common ancestor about
0\DZLWKFRQ¿GHQFHLQWHUYDOEHWZHHQDQG
0\D8SKDPDQG3DWWHUVRQ&DVWLOORHWDO
ZHUHWKH¿UVWWRSURYLGHHVWLPDWHVRISRWHQWLDO
ages of species of Ctenomys based on molecular clocks
RIPXOWLSOHJHQHWLFORFLLQWHUHVWLQJO\DQH[SDQGHGcytb
GDWDVHWDQDO\]HGE\WKRVHDXWKRUVVXJJHVWHGDQRULJLQ
for the species of CtenomysDWDERXW0\D7KLV
date is close to the estimate based on mitochondrial and
nuclear loci provided by Upham and Patterson (2012)
DWFD0\D,QFRPELQDWLRQWKHVHGDWDLQGLFDWHWKDW
the origin of species attributable to the genus Ctenomys
is likely to have occurred after the end of the rapid
RURJHQ\RIWKH$QGHV*DU]LRQHHWDO
$OOWKUHHNQRZQORFDOLWLHVIRUC. erikacuellarae
RFFXS\WKHVDPHULYHUGUDLQDJHVHH)LJ)LQRQOLQH
supplemental documents) and are not separated by
high elevation ridges, but geographic distance appears
important in differentiation among populations of
this species. Collection localities “Monteagudo” and
“Chuhuayaco” are separated by only about 16 km, (see
)LJ)LQRQOLQHVXSSOHPHQWDOGRFXPHQWVDQGWKHVH
two populations of C. erikacuellarae have a mean ge
netic distance of about 2.4% while comparisons of the
populations of Monteagudo and Vallegrande (separated
E\DSSUR[NPGLYHUJHE\Ctenomys andersoni shares a common ancestor with C. erikacuellarae
VHH)LJV)DQG)LQRQOLQHVXSSOHPHQWDOGRFX
ments) and genetic distance between these two species
LV*HRJUDSKLFGLVWDQFHDPRQJORFDOLWLHVRIWKHVH
two species crosses several major river valleys and
ULGJHVEXWVWUDLJKWOLQHGLVWDQFHUDQJHVIURPRQO\
NPVHH)LJV)DQG)LQRQOLQHVXSSOHPHQWDOGRFX
ments). Thus, alpha diversity in Ctenomys in Bolivia
DSSHDUVWREHLQÀXHQFHGE\WKHFRPSOH[JHRORJLFDQG
JHRJUDSKLFGLYHUVLW\RIWKHHDVWHUQVORSHRIWKH$QGHV
5HODWLYHWRELRGLYHUVLW\LQWKH<XQJDVDQG'U\
Valleys of Bolivia, we urge that additional broad scale
biological surveys of these areas be conducted before
anthropogenic impacts to native species are too severe.
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HJ *DUGQHU DQG &DPSEHOO D *DUGQHU DQG
BG$
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6DOD]DU%UDYR DQG<DWHV *DUGQHU HW DO
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discovery in this area. This strengthens the hypothesis
WKDWWKHFHQWUDO$QGHVEDFNWKUXVWEHOWLHWKHFRQWLQXDO
IRUPDWLRQ RI QHZ QRUWKVRXWK YDOOH\V DQG ULGJHV LQ
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Bolivia can be considered as a “speciation engine”
IRUELRGLYHUVLW\&KDUDFWHUL]LQJWKHH[WHQWRIQDWXUDO
diversity in this, or any geographic area, is a necessary
¿UVWVWHSWRZDUGHIIHFWLYHFRQVHUYDWLRQRILUUHSODFHDEOH
biotic resources.
-'H!)#,"5?8"!6%
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earnest in 1984 when a series of grants funded by the
National Science Foundation were awarded to Sydney
$QGHUVRQ7HUU\/<DWHV6FRWW/*DUGQHU-RVHSK$
&RRNDQG-RUJH6DOD]DU%UDYR%65WR6/*
%65WR6/*'(%WR6/*'(%
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6$ DQG -$& ,17 WR 7/< DQG -6% DQG
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WR6/*&ROODERUDWLQJLQVWLWXWLRQVLQ%ROLYLD
LQFOXGHG WKH 8QLYHUVLGDG$XWyQRPD *DEULHO 5HQp
0RUHQR WKH =RROyJLFR 0XQLFLSDO )DXQD 6XGDPHUL
FDQDERWKRI6DQWD&UX]WKH&ROHFFLyQ%ROLYLDQDGH
Fauna of the National Museum of Natural History, La
3D]DQGWKH%ROLYLDQ$FDGHP\RI6FLHQFHV/D3D]
1RHO.HPSII0HUFDGRODWHGLUHFWRURIWKH=RROyJLFR
Municipal Fauna Sudamericana, was instrumental in
DVVLVWLQJRXUZRUNLQWKHORZODQGVLQWKHPLGV
7LP.LOOHHQRIWKH0LVVRXUL%RWDQLFDO*DUGHQSURYLGHG
DVVLVWDQFHLQWKHVDQGWKHODWH*DVWRQ%HMDUDQR
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1980s.
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Fouisseur du Brésil. Nouveau Bulletin des Sciences
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62–64.
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of Ctenomys bicolor Miranda Ribeiro, 1914 (Ro
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16:445–447.
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72:56–61.
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uplift on climate and paleoaltimetry estimates.
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Felsenstein, J. 1985. Phylogenies and the comparative
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on chromosomal evolution in the neotropical cric
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ranean rodents of the genus Ctenomys (Rodentia:
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the Neotropics: Temporal and evolutionary impli
cations. Zoological Journal of the Linnean Society
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Bolivian mammals, 6: the genus Ctenomys (Ro
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Smithsonian Institution Press, Washington, D.C.
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of Linstowia =VFKRNNH &HVWRGD$QRSOR
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Baldellon. 2002. Tertiary provenance history of
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dentia: Octodontidae). Journal of Mammalogy
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Molecular Phylogenetics and Evolution 19:88–99.
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Addresses of authors:
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Harold W. Manter Laboratory
University of Nebraska State Museum and School of
Biological Sciences
University of Nebraska-Lincoln
Lincoln, Nebraska, USA 68588-0547
slg@unl.edu
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Department of Biological Sciences
Texas Tech University
Lubbock, Texas, USA 79409
j.salazar-bravo@ttu.edu
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Department of Biology and Museum of Southwestern
Biology
University of New Mexico
Albuquerque, New Mexico, USA 87131
tucojoe@gmail.com