Inglés

ISSN 0717-652X
Gayana 73(2): 161 - 171, 2009
REVALIDATION OF LATRODECTUS THORACICUS NICOLET, 1849
(ARANEAE: THERIDIIDAE): BIOLOGICAL AND PHYLOGENETIC
ANTECEDENTS
REVALIDACION DE LATRODECTUS THORACICUS NICOLET,
1849 (ARANEAE: THERIDIIDAE): ANTECEDENTES BIOLOGICOS Y
FILOGENETICOS
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ABSTRACT
The number and identity of the species of black widow spiders of the genus Latrodectus that inhabit Chile remain
XQFOHDU+HUHZHSUHVHQWUHVXOWVRIWD[RQRPLFZRUNEDVHGRQPRUSKRORJLFDQGPROHFXODUFKDUDFWHUVRIDODUJHVHULHVRI
specimens collected throughout Chile, which aimed to assess Chilean LatrodectusGLYHUVLW\5HVXOWVVKRZWKDWDOOVWXGLHG
populations belong to a single species, for which the correct name is Latrodectus thoracicus1LFROHW7KHUHIRUH
ZHUHPRYHWKLVWD[RQIURPWKHV\QRQ\P\RIL. mactans )DEULFLXV,QDGGLWLRQZHVXJJHVWWKDWL. mirabilis
(Holmberg, 1876) may be junior synonym of L. thoracicus:HUHFRJQL]HWZRVSHFLHVRIEODFNZLGRZVIRUFRQWLQHQWDO
Chile, L. thoracicus and L. variegatus1LFROHW:HSURYLGHGDQHPHQGHGGLDJQRVLVDQGFRPSUHKHQVLYHGHVFULSWLRQ
of L. thoracicusDVZHOODVQHZORFDOLW\UHFRUGVDQGQRYHOGDWDRQLWVQDWXUDOKLVWRU\*LYHQRXUUHVXOWVZHUHFRPPHQG
WKHFRQWLQXHGFROOHFWLQJVSHFLPHQVDQGPRUHFROOHFWLRQEDVHGWD[RQRPLFZRUNDVWKHRQO\ZD\RIJDWKHULQJDFRUUHFW
appreciation of LatrodectusGLYHUVLW\LQVRXWKHUQ6RXWK$PHULFD
KEYWORDS&KLOHVSHFLHVOLPLWVWD[RQRP\%ODFNZLGRZV
RESUMEN
(OQ~PHURHLGHQWLGDGGHODVHVSHFLHVGHYLXGDVQHJUDVGHOJpQHURLatrodectusTXHKDELWDQ&KLOHQRHVFODUR(QHVWH
WUDEDMRSUHVHQWDPRVUHVXOWDGRVGHXQHVWXGLRWD[RQyPLFREDVDGRHQFDUDFWHUHVPRUIROyJLFRV\PROHFXODUHVGHXQDVHULH
grande de especímenes colectados a lo largo de Chile, cuyo objetivo fue evaluar la diversidad de LatrodectusHQ&KLOH
Los resultados indican que las poblaciones estudiadas pertenecen a una misma especie, para la cual el nombre correcto
es Latrodectus thoracicus1LFROHW3RUORWDQWRUHPRYLPRVGLFKRWD[yQGHODVLQRQLPLDGHL. mactans (Fabricius,
$GHPiVVXJHULPRVTXHL. mirabilis (Holmberg, 1876) puede constituir un sinónimo junior de L. thoracicus
Reconocemos dos especies de viudas negras para Chile continental, thoracicus y L. variegatus1LFROHW3URYHHPRV
una diagnosis enmendada y una descripción detallada de L. thoracicus así como nuevas localidades de registro y datos
VREUHVXKLVWRULDQDWXUDO$ODOX]GHQXHVWURVUHVXOWDGRVD¿UPDPRVTXHHVLPSHULRVRFRQWLQXDUUHDOL]DQGRFROHFWDGH
HVSHFtPHQHV\WUDEDMRWD[RQyPLFREDVDGRHQFROHFFLRQHVFRPROD~QLFDIRUPDGHREWHQHUXQDFRUUHFWDDSUHFLDFLyQGH
la diversidad de LatrodectusHQHOVXUGH$PpULFDGHO6XU
PALABRAS CLAVE&KLOHOtPLWHVGHHVSHFLHVWD[RQRPtDYLXGDVQHJUDV
161
Gayana 73(2), 2009
INTRODUCTION
Latrodectus :DOFNHQDHU LV D GLYHUVH DQG
cosmopolitan genus of theridiid spiders with 30
UHFRJQL]HGVSHFLHV3ODWQLFN,WLVFRQVLGHUHGD
WD[RQRPLFDOO\FRPSOH[JHQXVDVWKHVWDWXVRIVHYHUDO
IRUPVKDYHQRWEHHQSURSHUO\HYDOXDWHGDQGVSHFL¿F
ERXQGDULHVDUHQRWZHOOGH¿QHGRUXQGHUVWRRG*DUE
et al/HYLDWKHUHIRUHLQPXOWLSOH
cases, populations have been uncritically referred to
DVGLIIHUHQWWD[D
The members of this genus are of medical importance
due to the potency of their venom, which contains
D QHXURWR[LQ WKDW SURGXFHV D PDVVLYH OLEHUDWLRQ
RI QHXURWUDQVPLWWHUV $PRQJ WKH SURWHLQV WKDW
cause the latrodectism one of the most important
is the DODWURWR[LQ 2UL ,NHGD *ULVKLQ
2UORYD et al %LRFKHPLFDO VWXGLHV
have been done to identify the proteins present in
some Chilean LatrodectusVSHFLPHQV7KHUHVXOWV
have been coincident with those based on North
American and European species (Romero et al
0HGLFDO DQG UHODWHG OLWHUDWXUH KDV
associated the Chilean cases of latrodectism with
Latrodectus mactans (Fabricius, 1775) (Artaza et
al $UWD]D et al 6FKHQRQH &RUUHD
5RPHURet al.3DURGLet al. 2008) or with
L. curacaviensis (Muller, 1776) (Faúndez 2009)
However, the number and identity of the species of
Latrodectus inhabiting Chile, is for the moment not
clear (Canals et alWKHUHIRUHLWLVQRWSRVVLEOH
to determine which species of Latrodectus cause
ODWURGHFWLVPLQ&KLOH
The genus Latrodectus has a wide distribution in
Chile, ranging from the deserts of the north (20ºS)
to the Patagonian steppes near the Magellan Strait
DSSUR[LPDWHO\ ž6 7KH QXPEHU DQG LGHQWLW\
of the species of Latrodectus currently inhabiting
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FRPSOH[ WD[RQRPLF KLVWRU\ RI WKH GLYHUVH IRUPV
of Latrodectus, and to the lack of studies that have
LQFOXGHG &KLOHDQ VSHFLPHQV 7KH ¿UVW UHFRUGV RI
Latrodectus in Chile were provided by Nicolet
(1849), who described Latrodectus variegatus
Nicolet 1849 and Latrodectus thoracicus Nicolet,
ZLWK,VOD*UDQGHGH&KLORpDQG&RQFHSFLyQ
DV WKH W\SH ORFDOLWLHV UHVSHFWLYHO\$W WKH VDPH
time, Nicolet (1849) mentioned a third form,
Latrodectus formidabilis :DOFNHQDHU IURP
&KLOHWKLVWD[RQZDVODWHUFRQVLGHUHGDVDV\QRQ\P
of L. mactansE\*HUVFKPDQ6FKLDSHOOL
162
and of L. curacaviensisE\/HYL+RZHYHU
Platnick (2009) points out that L. mactans is
not present in Chile and lists three Latrodectus
species for Chile: 1) Latrodectus geometricus &
/ .RFK ZLWK D ZRUOGZLGH GLVWULEXWLRQ Latrodectus curacaviensis (Müller, 1776) found in
WKH$QWLOODVDQG6RXWK$PHULFDZLWKWKHH[FHSWLRQ
RI$UJHQWLQDDQGL. variegatus Nicolet, 1849 with
a restricted distribution in Chile and adjacent zones
RI$UJHQWLQHDQDQG&KLOHDQ3DWDJRQLD1LFROHW
/HYLDE$EDORV,WVKRXOG
be noted that no revisionary work has included
more than one Chilean specimen of Latrodectus
/HYLLQFOXGHGRQHVSHFLPHQLGHQWL¿HGDVL.
curacaviensis7KH SK\ORJHQHWLF DQDO\VLV RI *DUE
et al LQFOXGHG D VLQJOH &KLOHDQ VSHFLPHQ
WKDWZDVQRWLGHQWL¿HGDWWKHVSHFLHVOHYHODQGIRU
which no information on its geographic origin was
SURYLGHG
The present study is based on the analysis of
morphological and molecular characters, and has
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distribution of the species of black widow present
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MATERIALS AND METHODS
MATERIAL
:HVWXGLHGDWRWDORIIHPDOHVPDOHVDQG
immature specimens from three Chilean collections:
Museo de Zoología de la Universidad de Concepción
(MZUC-UCCC), Museo Nacional de Historia
Natural (MNHN), and Laboratorio de Entomología
y Ecología, Universidad de La Serena (LEULS),
and the African National Museum Bloemfontein
10%$GGLWLRQDO VSLGHUV ZHUH FROOHFWHG E\ WKH
DXWKRUV XVLQJ D PDQXDO WHFKQLTXH 8SWRQ Sorensen et al IURP ORFDOLWLHV IURP
Arica and Parinacota Region in northern Chile to
the Magallanes and Antártica Chilena Region in
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MORPHOLOGICAL ANALYSIS
Morphological descriptions follow those of Lotz
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GHVFULSWLRQV DUH LQ PLOOLPHWUHV 7KH IROORZLQJ
WZR FRHI¿FLHQWV ZHUH FDOFXODWHG FHSKDORWKRUDFLF
FRHI¿FLHQW 7 FHSKDORWKRUD[ OHQJWK ZLWKRXW
FKHOLFHUDH GLYLGHG E\ FHSKDORWKRUD[ ZLGH 77
Revalidation of Latrodectus thoracicus: AGUILERA, M.A. ET AL.
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DQGWLELDRQOHJ,GLYLGHGE\FHSKDORWKRUD[OHQJWK
ZLWKRXWFKHOLFHUDH
PHYLOGENETIC ANALYSIS
Design and sampling 3K\ORJHQHWLF DQG JHQHWLF
analyses were based on a fragment of the
mitochondrial genome corresponding to the
F\WRFKURPHR[LGDVHVXEXQLW,&2,JHQH$QDO\VHV
LQFOXGHGVHTXHQFHVRIQLQH&KLOHDQVSHFLPHQVHLJKW
of these were collected by us at the following two
localities: 1) Casas Viejas, Comuna San Clemente,
5HJLyQ GHO 0DXOH ƒ¶´ 6 ƒ¶´
: Q DQG /RPDV /RV +XDVRV$QGDFROOR
5HJLyQ GH &RTXLPER ƒ¶´ 6 ƒ¶´ :
Q WKH QLQWK &KLOHDQ KDSORW\SH ZKLFK ODFNV
VSHFLHV GHWHUPLQDWLRQ DQG VSHFL¿F ORFDOLW\ ZDV
WDNHQIURP*HQ%DQN$<*DUEet al
The remaining sequences were downloaded from
*HQ%DQN7KXVRXUVDPSOLQJLQFOXGHVVHTXHQFHV
of Latrodectus IURP VSHFLHV 7DEOH $V WKH
monophyly of Latrodectus is well supported (Garb
et al., 2004) the outgroup was constituted by a single
representative of the false widow spiders theridiid
genus Steatoda
Sequence acquisition '1$ ZDV H[WUDFWHG XVLQJ
a commercial (Promega) kit from one or two
OHJV 7KH UHVW RI WKH VSHFLPHQV ZHUH SUHVHUYHG
DV YRXFKHUV$ IUDJPHQW RI WKH &2, JHQH ZDV
DPSOL¿HGZLWKSULPHUV/&2,DQG+&2
(Folmer et al$PSOL¿FDWLRQVZHUHGRQHLQ
three stages: 1) initial denaturation at 94 ºC for 90
VHFRQGVVF\FOHVRIGHQDWXUDWLRQDWž&
IRUVDQQHDOLQJDWž&IRUVDQGH[WHQVLRQ
DWž&IRUVDQG¿QDOH[WHQVLRQDWž&
IRUPLQ$PSOLFRQVZHUHVHQWWR0DFURJHQ,QF
.RUHDIRUSXUL¿FDWLRQDQGVHTXHQFLQJ2EWDLQHG
sequences were submitted to GenBank (accession
QXPEHUV*8
Sequence analysis6HTXHQFHDOLJQPHQWZDVGRQH
with Clustal X (Thompson et al XVLQJ
WKH GHIDXOW YDOXHV IRU DOO DOLJQPHQW SDUDPHWHUV
Observed percentage of sequence divergence was
calculated with MEGA 4 (Tamura et al. 2007)
LJQRULQJWKRVHVLWHVZLWKPLVVLQJGDWD*HQHWUHHV
ZHUHLQIHUUHGE\0D[LPXP3DUVLPRQ\)DUULV
using PAUP* (Swofford, 2000) with characters
WUHDWHGDVXQRUGHUHGDQGZLWKQRZHLJKWV7KHVHDUFK
strategy consisted of 200 replicates of heuristic
search with random addition of sequences and
7%5EUDQFKVZDSSLQJ6XSSRUWIRUWKHQRGHVZDV
DVVHVVHGE\D-DFNNQLIH-.DQDO\VLVFRQVLVWLQJRI
1000 pseudo-replicates with 2 replicates of random
addition of sequence, TBR branch swapping, and
RIFKDUDFWHUGHOHWLRQ*LYHQWKDWDQKDSORW\SH
was shared among newly sequenced specimens,
UHGXQGDQW VHTXHQFHV ZHUH H[FOXGHG IURP WKH
DQDO\VLVVHH)LJ
RESULTS
Of 646 characters, 186 were variable and 143 were
SDUVLPRQ\LQIRUPDWLYH7KHSK\ORJHQHWLFDQDO\VLV
UHFRYHUHGHTXDOO\SDUVLPRQLRXVWUHHVOHQJWK VWHSV&, 5, WKHFRQVHQVXVRI
ZKLFK)LJLVPRGHUDWHO\UHVROYHG$VLQ*DUEet
alWKHEDVDOGLFKRWRP\RIWKHLatrodectus
clade was the geometricus JURXS -. KHUH
represented by L. geometricus and L. rhodesiensis,
and the mactansJURXS-. ZKLFKFRQWDLQHG
the rest of the Latrodectus species included in our
VWXG\7KHUHZDVDZHOOVXSSRUWHGFODGH-. that contained Chilean haplotypes gathered by us,
haplotypes gathered by Garb et alIURPDQ
XQLGHQWL¿HG &KLOHDQ VSHFLPHQ RWKHU VSHFLPHQV
LGHQWL¿HG DV L. variegatus from the Argentinean
provinces of Río Negro and Santa Cruz, and a
KDSORW\SHUHFRYHUHGIURPDVSHFLPHQLGHQWL¿HGDV
of L. mirabilisIURP%XHQRV$LUHVSURYLQFH6LVWHUWR
this clade is a second haplotype of L. mirabilis also
from Buenos Aires Province, Argentina, which on
DYHUDJHGLYHUJHVE\IURPWKHFODGHFRQWDLQLQJ
WKH&KLOHDQKDSORW\SHVWKHUHVXOWLQJFODGHLVDOVR
ZHOOVXSSRUWHG-. Morphological (see below) and phylogenetic
analyses shown that all Chilean specimens analyzed
EHORQJ WR D VLQJOH VSHFLHV &RPSDULVRQV ZLWK WKH
GHVFULSWLRQVDQGPXVHXPVHULHVRIRWKHUWD[DDOORZ
us to conclude that the correct name to apply to
the studied populations is Latrodectus thoracicus
1LFROHWZKRVHUHGHVFULSWLRQIROORZV
Latrodectus thoracicus Nicolet, 1849
)LJV
Latrodectus thoracicus1LFROHW
Latrodectus formidabilis1LFROHWODP
I
Latrodectus mactans. *HUVFKPDQ 6FKLDSHOOL
Latrodectus curacaviensis/HYL
163
Gayana 73(2), 2009
TABLE I /LVWRIVSHFLPHQVIURPZKHUHKDSORW\SHVRIWKH&2,JHQHXVHGLQWKHSK\ORJHQHWLFDQDO\VLVZHUHJDWKHUHG
For those sequences generated here numbers of corresponding voucher specimens are provided (for locality data see
0DWHULDO([DPLQHG6HTXHQFHPDUNHGZLWKDQDVWHULVNZDVIURPDQXQLGHQWL¿HG&KLOHDQVSHFLPHQDQGKHUHUHIHUUHG
to L. thoracicus
TABLA I /LVWDGH HVSHFtPHQHVDSDUWLUGHORVFXDOHVVHJHQHUDURQORVKDSORWLSRVGHOJHQ&2,XVDGRVHQHODQiOLVLV
¿ORJHQpWLFR3DUDDTXHOODVVHFXHQFLDVJHQHUDGDVHQHVWHHVWXGLRWDPELpQVHLQGLFDORVFRUUHVSRQGLHQWHVQ~PHURVGHORV
HVSHFLPHQHVYRXFKHUVSRUGDWRVGHORFDOLGDGHVYHU0DWHULDO([DPLQHG(ODVWHULVFRLQGLFDXQDVHFXHQFLDJHQHUDGDSRU
Garb et alDSDUWLUGHXQHVSpFLPHQFKLOHQRQRLGHQWL¿FDGR\TXHDTXtHVUHIHULGRDL. thoracicus
species
specimen voucher
accession number
source
Ingroup
1
L. antheratus
AY383048
Garb et al. 2004
2
L. antheratus
AY383047
Garb et al. 2004
3
L. corallinus
AY383062
Garb et al. 2004
4
L. corallinus
AY383061
Garb et al. 2004
5
L. diaguita
AY383063
Garb et al. 2004
6
L. diaguita
AY383064
Garb et al. 2004
7
L. geometricus
AY383066
Garb et al. 2004
8
L. hasselti
EF121035
Vink et al
9
L. hesperus
AY383071
Garb et al. 2004
10
L. katipo
EF121016
Vink et al
11
L. mactans
AY383072
Garb et al. 2004
12
L. menavodi
AY383075
Garb et al. 2004
13
L. mirabilis
AY383076
Garb et al. 2004
14
L. mirabilis
AY383077
Garb et al. 2004
15
L. pallidus
AY383055
Garb et al. 2004
16
L. rhodesiensis
AY383079
Garb et al. 2004
17
L. thoracicus
AY383074*
Garb et al. 2004
18
L. thoracicus
LT0204Cv01
GU112098
this study
19
L. thoracicus
LT0105Cv01
GU112099
this study
20
L. thoracicus
LT0105Cv02
GU112100
this study
21
L. thoracicus
LT0105Cv03
GU112101
this study
22
L. thoracicus
LT0105Cv05
GU112102
this study
23
L. thoracicus
LT0108Cv01
GU112103
this study
24
L. thoracicus
LT0108Cv02
GU112104
this study
25
L. thoracicus
LT0903An01
GU112105
this study
26
L. tredecimguttatus
AY383080
Garb et al. 2004
27
L. variegatus
AY383084
Garb et al. 2004
28
L. variegatus
AY383083
Garb et al. 2004
29
L. variolus
AY383082
Garb et al. 2004
AY383086
Garb et al. 2004
Outgroup
30
164
Steatoda grossa
Revalidation of Latrodectus thoracicus: AGUILERA, M.A. ET AL.
FIGURE 16WULFWFRQVHQVXVWUHHRIHTXDOO\SDUVLPRQLRXVWUHHVOHQJWK VWHSV&, 5, UHVXOWLQJ
from the analysis of haplotypes of the COI gene of Latrodectus6SHFLPHQVRIL. thoracicus LT0204Cv01, LT0108Cv01,
DQG/7&YVKDUHWKHVDPHKDSORW\SHZLWKVSHFLPHQ/7&YZKLFKLVPDUNHGZLWKDQDVWHULVN7KLVKDSORW\SH
LVDOVRVKDUHGE\DQXQLGHQWL¿HG&KLOHDQVSHFLPHQ$<VHTXHQFHGE\*DUEet aO
FIGURA 1&RQVHQVRHVWULFWRGHiUEROHVPiVFRUWRVORQJLWXG SDVRV,& ,5 UHVXOWDGRVGHO
análisis de haplotipos del gen COI de Latrodectus/RVHVSHFtPHQHVGHL. thoracicus LT0204Cv01, LT0108Cv01 y
/7&YFRPSDUWHQHOKDSORWLSRFRQHOHVSpFLPHQ/7&YHOFXDOVHLGHQWL¿FDFRQXQDVWHULVFR(VWHKDSORWLSR
HVWDPELpQFRPSDUWLGRSRUXQHVSpFLPHQFKLOHQRVLQLGHQWL¿FDU$<VHFXHQFLDGRSRU*DUEet al
165
Gayana 73(2), 2009
HOLOTYPE
$IHPDOHIURP3URYLQFLDGH&RQFHSFLRQ&KLOH7KH
W\SHLVORVW
EMENDED DIAGNOSES
Opening of the epigynum oval to suboval, with
anterior and posterior lips similar in length, and
anterior margin often with a small lobulated median
SRLQWHGSURFHVV)LJD&RQQHFWLQJGXFWVRIWKH
VSHUPDWKHFDZLWKWKUHHORRSVLQGRUVDOYLHZWKHVHFRQG
and third ones located at the side of the spermatheca and
URWDWHGRYHUWKHPVHOYHVLQDVLQJOHSODQHWKHODVWORRS
RXWOLQHDOPRVWDQHQWLUHFLUFOH)LJFG(PEROXVRI
WKHPDOHSDOSXVZLWKWKUHHFRLOV)LJDWKHFRQGXFWRU
placed in an angle larger than 90º, stout and strong in the
GLVWDOWKLUG)LJFUDGL[ODQFHRODWH)LJG
FIGURE 3: Latrodectus thoracicus: a) ventral view of the
epiginium, b) dorsal view of the epiginium, c) ventral
view of the internal genitalia, d) dorsal view of the internal
JHQLWDOLDDQGHODWHUDOYLHZRIWKHOHIWVSHUPDWHFD
FIGURA 3: Latrodectus thoracicus: a) vista ventral del
HSLJLQLREYLVWDGRUVDOGHOHSLJLQLRFYLVWDYHQWUDOGH
la genitalia interna, d) vista dorsal de la genitalia interna,
\HYLVWDYHQWUDOGHHVSHUPDWHFDL]TXLHUGD
166
DESCRIPTION:
Female&HSKDORWKRUD[OHJVDQGDEGRPHQDUHVKLQ\
EODFN)LJE$EGRPHQZLWKGRUVDOUHGRURUDQJH
markings that are diverse in shape and arrangement
)LJFGYHQWUDOO\ZLWKDWUDSH]RLGPDUNZLWK
a median cut between the spinnerets and epigastral
IXUURZWKHVKDSHRIWKLVPDUNLVVOLJKWO\YDULDEOH
EXWLVJHQHUDOO\WKHFODVVLFKRXUJODVVVKDSH)LJH
Pair of circular spots present on each side of the
ODWHUDODQWHULRUDQGODWHUDOSRVWHULRUVSLQQHUHWV)LJ
I7KHSDWWHUQRIWKHVHVSRWVYDULHVDFFRUGLQJWR
DJHDQGJHRJUDSK\)LJFGI7KHDEGRPLQDO
FIGURE 4: Latrodectus thoracicus: a) ventral view of the
left palpus, b) ectal view of the left palpus, c) right palpus
FRQGXFWRUGULJKWSDOSXVUDGL[HYHQWUDOYLHZRIWKH
ULJKWSDOSXVDQGILPPDWXUHVSHFLPHQ< F\PELXP
3 SDUDF\PELDO KRRN 6 VXEWHJXOXP 7 WHJXOXP
0 PHGLDQ DSRSK\VLV ( HPEROXV 5 UDGL[ & FRQGXFWRU
FIGURA 4: Latrodectus thoracicus: a) vista ventral del
SDOSR L]TXLHUGR E YLVWD HFWDO GHO SDOSR L]TXLHUGR F
FRQGXFWRUGHOSDOSRGHUHFKRGUDGL[GHOSDOSRGHUHFKR
HYLVWDYHQWUDOGHOSDOSRGHUHFKR\IHVSpFLPHQMXYHQLO
<&LPELR3*DQFKRGHOFLPELR6VXEWHJXOR7WHJXOR
0DSy¿VLVPHGLD(HPEROR5UDGL[&FRQGXFWRU
Revalidation of Latrodectus thoracicus: AGUILERA, M.A. ET AL.
77FRHI¿FLHQW
MaleVPDOOHUWKDQWKHIHPDOH&HSKDORWKRUD[GDUN
EURZQZLGHUWKDQORQJHU$EGRPHQRIIZKLWHZLWK
IRXUGDUNGRUVDOEDQGVWZRRIWKHPDUHLUUHJXODU
and located in the antero-medial part and project
WRWKHVLGHVWKHRWKHUWZREDQGVDUHHORQJDWHGDQG
GLUHFWHGWRWKHSRVWHULRUHQG/HJVORQJHUWKDQLQ
WKHIHPDOHFRORXUGDUNEURZQZLWKFUHDP\ZKLWH
VWULSHV)LJD0DOHSDOSDQGEXOEW\SLFDOIRUWKH
JHQXVHPERORXVZLWKWKUHHFRLOV)LJDEH
the conductor is placed in an angle larger than 90º,
VWRXWDQGVWURQJLQWKHGLVWDOWKLUG)LJFUDGL[
ODQFHRODWH )LJ G 0HDVXUHPHQWV DV IROORZV
Q WRWDOOHQJWKFHSKDORWKRUD[
OHQJWK ZLGWK 7FRHI¿FLHQWOHQJWKWLELD,
OHQJWKSDWHOOD,77
FRHI¿FLHQW
FIGURE 2: Latrodectus thoracicus: a) adult male, b-f) adult
female, c-d) variations of dorsal pattern coloration, e)
YHQWUDOSDWWHUQDQGISDWWHUQDURXQGVSLQQHUHWV
FIGURA 2: Latrodectus thoracicus: a) macho adulto, b-f),
hembra adulta, c-d) variaciones de patrones de manchas
dorsales, e) patrón ventral, y f) manchas alrededor de las
hilanderas.
GRUVXPZLWK¿QHFXUYHGVHWDHRIGLIIHUHQWOHQJWKV
(SLJ\QXPVXEJORERLGUXJRVHDQGSURMHFWLQJZLWK
an opening oval to suboval, wider than longer, with
sclerotized lips and the anterior margin often with a
small median pointed process as a lobule, and both
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connecting ducts with three loops, the second and
third ones located at the side of the spermatheca and
rotated over themselves in a single plane, while the
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Immatures: Around 55 days after emergence,
individuals show basic characters of the genus
(serrate setae on metatarsus IV, evident colulus with
two setae, lateral eyes apart, and lack of teeth of the
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WKH FHSKDORWRUD[ DQG OHJV EURZQRUDQJH DQG
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of the abdomen with a series of rhomboidal dark
EURZQ VSRWV )LJ I 9HQWHU ZLWK IRXU FLUFXODU
and yellowish spots located around the spinnerets,
and with a yellowish outline of the adult trapezoid
spot located between the epigastral furrow and the
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Eggs: 6SKHULFDO\HOORZLVKOHQJWKEHWZHHQDQG
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egg sac is 536 rQ Egg-sac Q VSKHURLG RU VRPHWLPHV SHDU
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COMPARISONS
Females of L. thoracicus differ from those of the
other Latrodectus species mentioned for Chile, L.
variegatus, L curacaviensis, and L. geometricus
(Platnick 2009), by the arrangement of the
spermathecae, the number of loops of the genitalia
167
Gayana 73(2), 2009
ducts, and the arrangement of the dorsal spots of
WKHDEGRPHQLatrodectus geometricus has parallel
spermathecae while those of L. thoracicus are placed
LQ D 9IRUPDWLRQ Latrodectus variegatus and L.
curacaviensis (Abalos 1980) have spermathecal
ducts with two loops, and L. thoracicus has three
ORRSVLatrodectus curacaviensis has also a black
abdomen with an elongated central band, surrounded
by three pairs of bands placed obliquely (fide
0F&URQH/HYL$EDORVLatrodectus
thoracicus, on the other hand, has a black abdomen
but with spots or bands of different shapes and
YDULDEOHDUUDQJHG
Males of L. thoracicus differ from those of L.
variegatus, L. geometricus, and L. curacaviensis by
the number of loops of the embolus and the shape
RIWKHFRQGXFWRU7KHHPEROXVRIL. variegatus has
two loops, that of L. geometricus has four, while that
of L. thoracicusKDVWKUHHORRSV,QL. curacaviensis
the distal third of the conductor is more or less
cylindrical and in L. thoracicusLVGLODWHG
DISTRIBUTION
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from Arica y Parinacota Region in the north (20°S)
to Barranca Negra in Magallanes Region in the
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DUHDVRI$UJHQWLQHDQ3DWDJRQLD,WLVDEXQGDQWLQWKH
FHQWUDODQGVRXWKHUQSDUWVRI&KLOH
MATERIAL EXAMINED
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168
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Revalidation of Latrodectus thoracicus: AGUILERA, M.A. ET AL.
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NATURAL HISTORY
In the Central Valley of Chile L. thoracicus can be
found in sunny areas with or without low herbaceous
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ground, under pieces of wood and rocks or at the
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associated with agricultural plantations such as
those of oat, corn, alfalfa, and vineyards, where
they are located close to the ground, under bales,
among leaves and trunks of vineyards, blackberries
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The webs of L. thoracicus are irregular and are
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SRVVLEOH WR GLVWLQJXLVK VRPH VSHFL¿F DUHDV LQ WKH
ZHE¶V VWUXFWXUH UHIXJH ]RQH ZKHUH WKH VSLGHU
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ZDVWH]RQHZKHUHWKHVSLGHUOHDYHVWKHIRRGUHVWV
capture zones, which correspond to all web places
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zone can be seen in the web where undigested preys
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ground are more or less square and can be from
IHZFHQWLPHWUHVWRXSPHWHUVZLGH+RZHYHU
ZHEVEXLOWRQYHJHWDWLRQFDQH[WHQGHGORQJHUWKDQ
2 meters from bottom to top and intermingle with
WKHYHJHWDWLRQ
The most common prey of L. thoracicus are
locusts, crickets, tenebrionids, scarabaeids, isopods,
JUDVVKRSSHUVDQGDOVRRWKHUVSLGHUV2FFDVLRQDOO\
L. thoracicusIHHGVRQZDVSV
Specimens can be found throughout most of the
\HDUEXWWKHSRSXODWLRQÀXFWXDWHVGXULQJWKH\HDU,Q
the 2008 austral summer, two males were collected
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trunks in a vineyard at San Clemente, Región del
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DISCUSSION
Latrodectus thoracicus was described by Nicolet
(1849) on the basis of one specimen collected
QHDU&RQFHSFLyQFHQWUDO&KLOH,WZDVFRQVLGHUHG
as a junior synonym of L. mactans by Gertchman
6FKLDSHOOL DQG ODWHU RI Latrodectus
curacaviensis E\ /HYL +RZHYHU LQ WKHVH
last two papers neither type nor topotypical material
ZDVH[DPLQHG7KHFKDUDFWHUVPHQWLRQHGE\1LFROHW
(1849) in his description of L. thoracicus match
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7KHUHIRUH ZH VXJJHVW WKDW 1LFROHW¶V GHVFULSWLRQ
was based on an immature specimen and that the
specimens here studied belong to L. thoracicus
&XUUHQWO\DIWHU*HUVFKPDQ6FKLDSHOOLL.
thoracicus is considered as a junior synonym of L.
mactansDQGDVVXFKLVOLVWHGVSHOOHGDVL. thoracius)
in Platnick`s (2009) catalogue, even when he does
not include Chile in the distribution of L. mactans
Our assessment of patterns of morphological
variation and the results of the phylogenetic analysis
clearly indicate that L. thoracicus differs from
both L. mactans and L. curacaviensis 6LPLODUO\
L. thoracicus morphologically differs from L.
variegatus, another Chilean species described from
&KLORH,VODQG7KHUHIRUHZHFRQVLGHUL. thoracicus
LVDYDOLGVSHFLHV
Nicolet (1849) also mentioned L. formidabilis from
Chile, but based on the information provided in
his short description, we believe this record may
represent L. thoracicus
There is low genetic diversity of COI in populations
of L. thoracicus KDSORW\SHV UHFRYHUHG IURP
Chilean populations diverge on average by only
WKLV YDOXH LV LQ OLQH ZLWK WKRVH IRXQG E\
Vink et alIRUVSHFLHV IURP1HZ=HDODQG
The sample of L. thoracicus from San Clemente
VKRZV PLQLPDO KDSORW\SLF GLYHUVLW\ )RXU RI
seven specimens from this locality share the same
KDSORW\SH SDLUZLVH FRPSDULVRQV JLYH DQ DYHUDJH
YDOXH IRU WKLV VDPSOH RI 5HPDUNDEO\ WKH
169
Gayana 73(2), 2009
most abundant haplotype found in San Clemente
was also recovered from Argentinean specimens
LGHQWL¿HGE\*DUEet alDVEHORQJLQJWRL.
miriabilis and L. variegatus from Buenos Aires, Río
1HJURDQG6DQWD&UX]SURYLQFHV7KLVXQH[SHFWHG
similarity may be due to the short sequence lengths
(430 base pairs) obtained by Garb et alIURP
the Argentinean specimens and/or because species
have retained an ancestral genetic variant (Pamilo
1HL+RZHYHUWKLV¿QGLQJTXHVWLRQVWKH
distinction of these nominal forms or at least the
correct determination of the specimens from where
WKHVHKDSORW\SHVZHUHJDWKHUHG,WLVSRVVLEOHWKDWL.
miriabilis is a junior synonym of L. thoracicus:H
GLGQRWSURSRVHDQ\IRUPDOFKDQJHWRWKHWD[RQRPLF
status of L. miriabilis at this stage, because we have
QRWKDGWKHRSSRUWXQLW\WRH[DPLQHWKHVSHFLPHQV
sequences by Garb et alWRFKHFNWKHLUFRUUHFW
LGHQWLW\Latrodectus thoracicus is morphologically
distinguishable from L. variegatus as described
E\1LFROHW7KHUHIRUHZHVXJJHVWWKDWWKH
VSHFLPHQV LGHQWL¿HG E\ *DUE et al DV L.
variegatus from Argentina belong in fact to L.
thoracicus
Two species of Latrodectus inhabit continental
&KLOH 2QH VSHFLHV L. variegatus, seems to be
UHVWULFWHGWR&KLORH,VODQG7KHVHFRQGVSHFLHVL.
thoracicus LV ZLGHO\ GLVWULEXWHG LQ WKH FRXQWU\
:H VXJJHVW WKDW IXWXUH VWXGLHV VKRXOG LQFOXGH
the analysis of type material as well as of newly
collected specimens from type localities as the
RQO\ZD\RIDVVLJQLQJWKHFRUUHFWWD[RQRPLFQDPHV
WR PLWRFKRQGULDO KDSORW\SHV DQG PRUSKRW\SHV
:KDWHYHU WKH UHVXOWV RI WKHVH VWXGLHV PLJKW EH
LW LV VDIH WR DVVXPH WKDW WKH FXUUHQW WD[RQRPLF
scenario does not correctly reflect the alpha
diversity of the genus Latrodectus and/or that
VSHFLHV¶ JHRJUDSKLF UDQJHV DUH SRRUO\ GHOLPLWHG
Given the medical relevance of this genus, it
ZRXOG EH YDOXDEOH WR FRQGXFW IXUWKHU ¿HOG ZRUN
DQGFROOHFWLRQEDVHGWD[RQRPLFZRUNLQRUGHUWR
gain a correct appreciation of Latrodectus diversity
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170
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REFERENCES
ABALOS, J.W /DV DUDxDV GHO JpQHUR Latrodectus
HQOD$UJHQWLQD2EUD&HQWHQDULDGHO0XVHRGH
/D3ODWD
ARTAZA O., FUENTES J. & SCHINDLER, R/DWURGHFWLVPR
HYDOXDFLyQ FOtQLFRWHUDSpXWLFD GH FDVRV
5HYLVWD0pGLFDGH&KLOH
ARTAZA O., FUENTES J., GÓMEZ P. & MORRIS R /DWURGHFWLVPR,,HYDOXDFLyQFOtQLFRWHUDSpXWLFD
GHFDVRV3DUDVLWRORJtDDO'tD
CANALS, M., CASANUEVA, M.E. & AGUILERA M.A.
¿Cuáles son las especies de arañas peligrosas en
&KLOH"5HYLVWD0pGLFDGH&KLOH
FARRIS, J.S 7KH ORJLFDO EDVLV IRU SK\ORJHQHWLF
DQDO\VLV In 3ODWQLFN , )XQN 9$ (GV
$GYDQFHV LQ &ODGLVWLFV 3URFHGLQJV RI WKH
VHFRQGPHHWLQJ:LOOL+HQQLQJ6RFLHW\&ROXPELD
8QLYHUVLW\3UHVV1HZ<RUN9RO
F AÚNDEZ (, $UDxDV $UDFKQLGD $UDQHDH
SHOLJURVDV GH OD 5HJLyQ GH 0DJDOODQHV$QDOHV
Instituto Patagonia FOLMER, O., BLACK, M., HOEH, W., LUTZ, R. & VRIJENHOEK,
R., '1$ SULPHUV IRU DPSOLILFDWLRQ RI
PLWRFKRQGULDO F\WRFKURPH F R[LGDVH VXEXQLW ,
IURPGLYHUVHPHWD]RDQLQYHUWHEUDWHV0ROHFXODU
Marine Biology and Biotechnology 3(5):294
GARB, J. E., GONZÁLEZ, A. & GILLESPIE, R.G. 20047KH
black widow spider genus Latrodectus (Araneae:
Theridiidae): phylogeny, biogeography and
LQYDVLRQ KLVWRU\ 0ROHFXODU 3K\ORJHQHWLFV DQG
(YROXWLRQ
GERSCHMAN, B.S. & SCHIAPELLI R.D5HYLVLyQGHO
JpQHURLatrodectus:DOFNHQDHUIn Sampayo,
5Latrodectus mactans \ODWURGHFWLVPR%XHQRV
$LUHV6\VW
GRISHIN, E 3RO\SHSWLGH QHXURWR[LQV IURP VSLGHU
YHQRPV (XURSHDQ -RXUQDO RI %LRFKHPLVWU\
LEVI, H.W7KHVSLGHUJHQXVLatrodectus (Araneae,
7KHULGLLGDH 7UDQVDFWLRQV RI WKH $PHULFDQ
Microscopical Society 78:
LEVI, H.WD&RVPRSROLWDQDQGSDQWURSLFDOVSHFLHV
RIWKHULGLLGVSLGHUV$UDQHDH7KHULGLLGDH3DFL¿F
,QVHFWV
LEVI, H.WE7KH7KHULGLLGVSLGHUIDXQDRI&KLOH
Bulletin of the Museum of Comparative Zoology
Revalidation of Latrodectus thoracicus: AGUILERA, M.A. ET AL.
LOTZ, L.N 5HYLVLRQ RI WKH JHQXV Latrodectus
$UDQHDH 7KHULGLLGDH LQ$IULFD 1DYRUVLQJH
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ORI, M. & IKEDA, H 6SLGHU 9HQRPV DQG 6SLGHU
7R[LQV -RXUQDO RI 7R[LFRORJ\ 7R[LQ UHYLHZV
ORLOVA E.V., RAHMAN A., GOWEN B., VOLYNSKI K.E.,
A SHTON A.C., M ANSER C, VAN H EEL , M. &
USHKARYOV, Y.A. 6WUXFWXUHRIĮODWURWR[LQ
oligomers reveals that divalent cation-dependent
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PARODI, J., ROMERO, F., MILEDI, R. & MARTÍNEZ, A. Some effects of the venom of the Chilean spider
Latrodectus mactans on endogenous ion-currents
of Xenopus laevis RRF\WHV %LRFKHPLFDO DQG
Biophysical Research Communications 375(4):571
PLATNICK, N.I.7KH:RUOG6SLGHU&DWDORJ9HUVLRQ
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ROMERO, F., ALTIERI, E., QUIÑEHUAL, C. & CAYUQUEO, A.
$FWLYLGDG FRQWUiFWLO GHO P~VFXOR SDSLODU
cardiaco y conducto deferente de rata inducida
por veneno de la araña Latrodectus mactans de
&KLOH*D\DQD
ROMERO, F., ALTIERI, E., URRUTIA, M. & JARA, J. Venom of Latrodectus mactans from Chile
(Araneae, Theridiidae): effect on smooth muscle
5HYLVWDGH%LRORJtD7URSLFDO
ROMERO, F., SALVATICI, R., SCHULZ, C., CABEZAS, M.,
S TANDEN , D. & A LTIERI , E. (IHFWR GHO
veneno de Latrodectus mactans sobre la respuesta
FRQWUiFWLO GHO P~VFXOR SDSLODU GH UDWD 5HYLVWD
&KLOHQDGH&DUGLRORJtD
SCHENONE, H. & CORREA, L$OJXQRVFRQRFLPLHQWRV
prácticos sobre la biología de la araña Latrodectus
mactans\HOVtQGURPHGHODWURGHFWLVPRHQ&KLOH
%ROHWtQ&KLOHQRGH3DUDVLWRORJtD
SORENSEN, L., CODDINGTON, J.A. & SCHARFF, N Inventorying and estimating subcanopy spiders
diversity using semiquantitative sampling
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TAMURA, K., DUDLEY, J., NEI, M. & KUMAR, S.0(*$
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