Diversitat Del Gènere Loxosceles Heineken & Lowe, 1832 a La Mediterrània I Les Illes Canàries: Sistemàtica, Biogeografia I Loxoscelisme

Diversitat del gènere Loxosceles Heineken & Lowe, 1832 a la Mediterrània i les Illes Canàries: sistemàtica, biogeografia i loxoscelisme

Enric Planas Figueras

Aquesta tesi doctoral està subjecta a la llicència Reconeixement- NoComercial – CompartirIgual 3.0. Espanya de Creative Commons.

Esta tesis doctoral está sujeta a la licencia Reconocimiento - NoComercial – CompartirIgual 3.0. España de Creative Commons.

This doctoral thesis is licensed under the Creative Commons Attribution-NonCommercial- ShareAlike 3.0. Spain License. Diversitat del gènere Loxosceles Heineken & Lowe, 1832 a la Mediterrània i les Illes Canàries: sistemàtica, biogeografia i loxoscelisme

Enric Planas Figueras

TESI DOCTORAL Foto portada: Fuerteventura (E. Planas) Foto contraportada: Anti-Atles (E. Planas) Dibuix contraportada: Descripció de Loxosceles citigrada (= L. rufescens) a Heineken & Lowe, 1832. ! " # $!

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INTRODUCCIÓ

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4 INTRODUCCIÓ

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6 INTRODUCCIÓ

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8 INTRODUCCIÓ

C- D C 7118%9 U9 71P3E" ( #" " ,4, , /7 @ 71128C- D C 7118= 7113E " ,T ( - ,4 #, 4% T @,- ( " / RS 7173 $ $/%9 U9 71P3 6 "- ( - " , # ( , &" 2/0 711A3E" ,T, ,E75 ; ( /0 711A3% T , # $% /)&?78! - 0 711>3 S ( ($ ( " " ( 4 4 " /0 711A3 E # ($ ,( /7 713 " # , 4S 4" /0 711A3 , * +/= 7173 4" ." ."% " # 4 - /0 711P3 #, , 4/B@= @71123 ,- # 4 4 %S " N- ) - "4N " ,4 # S , 7 V ,T "<4 ,E75 -

9 6( &&$E" 4 # / @ ? 7136 " - &&$ - ,T & E" , #" ( , # /WDD = 23" ( # 4 ( ( , #/) - 711P35 ($ ,(4 , ( , # & ,- " , " * ( + C ,T = - 4 , 4 " $ $/) - =@711A80711>8 C- 71180 @ 7113 ? $&&$ , " # $/E71113 # - E" 4 /0D 7113 " . , 4/)%9 7113% , # , 4 4 ,E75 " / , # 4 # 8 E 71113 B " ,E75 ,( -4 S , ## 7 4 4 , T 4 - $ $&&$3/U9 B 711780D

10 INTRODUCCIÓ

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12 INTRODUCCIÓ

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13 711736 ( , #, " @ 4 & /) @ ) E713 C . -$ $ B , %5'" # (# . B ,E 2 B N -. B B /US D8 US 71178!-@' 7113C " $/) -B71113 $ , Z % T E" 2AAB " ,% O , ,E B B /O ' 7113 , # /= ' 71118 ' 7118C 71780 71780 71A 43

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14 INTRODUCCIÓ

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16 INTRODUCCIÓ

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18 INTRODUCCIÓ

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20 INTRODUCCIÓ

O- V /E 4 C' 53 " 4 /R)3

< =G< ?G = G G :H J :KHLM

? , 4 $ " , @ # # % S ; " 4 , @ " ( $ @ , @/071123 ($ @ - $ # " -

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22 INTRODUCCIÓ

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24 INTRODUCCIÓ

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25 - $ # 4 /C9 7118;7118' 7118O D 71P35-- ( # ( " - V , , , 4 ( , , @

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= " ( ,- 4 $ N ,4 " - @ " " ,- 4 $ ( ' " . (- , " # ( S $ S " . S - /; 71178 C9 711>8 $7 0 @7113E ,- " . / &. $ (

26 INTRODUCCIÓ

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OBJECTIUS

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32 OBJECTIUS

# # # $ # / # 5 # / > 7 )!

= >< C) 0 & ! # 0 # 0 & # 7 / # 5- & - / # # > !

= % >7 0 ) / 0 # # < - / 8 - # !

RESULTATS

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/ 6 / - - - / ? ' * 0 & - - ' /12/3 / 6 ? ' O+ L /3 / - 6 /

/ 6 / / ? - ' * 0 / - 6

38 RESULTATS

/ 6 / / ? -/ *L ' O 0 / - 6

/ 6 / / ? - 9 ? 9 / ? ' *' 0 / - 6

/ 6 / / ? - 9 ? /3G / ? ' / - 6

6 / ? / *L * / ? P - 6 /

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CAPÍTOL 1

Diversitat del gènere Loxosceles a la Conca Mediterrània

Article 1 Planas E, Saupe EE, Lima-Ribeiro MS, Peterson AT, Ribera C. Ecological niche and phylogeography elucidate complex biogeographic patterns in Loxosceles rufescens (Araneae, Sicariidae) in the Mediterranean Basin.

RESULTATS

! " # $ ! %" & ' ( # ! ) ! " * + ,, ( %,' # - + %.' # ! / 0 - 1 2 - * %34#'! & 5 ! 3 %34#' # "64 ! 7 4 8 1 - ! " 2 # 4 8 9 %":;#' * ! 3

43 Article 1

34# 2 < !

"#! #"( - 5 !

44 RESULTATS

! " #$

% " " &"' (%&) * "&+ ,' "& * -.+/ ,' /-0 ! 0 , &"' /% ,' /-0 ! 0 , * "$1 &2+ ,' " "3" ".4 $ 5 6 .7 &

," "+' / /-+ // 8 "-- " " ' ++ ! ++ " ' "-"/ " "" " & ! "-' +/-+ " ++ ""' / ( " ) " 8 9 ' " " "!"!"' ! ' "/' " " "' / ++ + / + + "" "++

* / / 8" % "-" "-- ' / + + ! +/++ 9( ) + ! ! " " % " () + :+ ++ " & * + / " "' - ' " -+ "-" + + "+ (;) ! "-- - + < " " ' " ++ /- " / " / "

=+ "+ " !/++ / "

45 Article 1

' / /- " & ! + -*; "' / =-" -/ - ! -+- " +"' /+ " / :/"/ '+ "+ = " - -+ ! " / - + -> + " ( 7.$)! !"---+" "+ ; ! --'"+ + '! - ' -+ ! - " ""+ ' - +/ "- + +"

!"#$%& " ' " " " + -+ "

!'$()%

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46 RESULTATS

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

47 Article 1

' / " "' /' -/++ " ; 7 !:+ / *; "' / ! + " /+ -+ " " ++ / ! - " / '/ / -- - +9( )"'+ ""!" - " + (FGHBB/) () '-+ " '8 "'" -+-" ' / + "()!" " + " ! !" &

*$%

= " " & " ' +(* B B) " :"- " '+! + % ?> ! " > (3+ " "" - )3 B"'" ! M" """ /

= "" "'" - / / + *; ! 8 " + * 77 *; 8 0 (&%7 77) -!+ - A = -" - / 8" %( ) +%;3 ". S ( # B ) "+ B G- + K! ""-- -$ 5 D > "$ 5 >( JJ) "$ ; J (3 JJ) ! - - "$ !"" - '-?T + / (+ )& *; / (& )$" ! / M"" + $ "&+*/ ' $/M+0( "& / )M !"'"- " &%DBB " M ' $6- U - ,' "& " + %( 0 ) ! M !"" " "!.;%7,G?(*" BBJ) '" - - "+ "/* (B B)- ! / " - - -- + K! "

48 RESULTATS

M " ":"- " M ! +" + /.;%7, +$ =( JJ)!"- = "" /" "8" + " " / + 3"' B ( - B )" & / - - 8 !""""/-"'" /"" / " -"" "" + /

k m k k mk m m km m k m m m k

+ = " 8:"() "& / -(&%)-/+ - " + '- / / !"" 8D( : BBG)! "-+ -" V.,% (' # : B )= " "+ + """ BBB &% / !""& / (M B )!!"" - + !- :' (" ") +'/ BBB + =:"'+- ' / ( #*" BBD) --' N () ! ' BB " ' + " " "' - -M ( *3)-! !!BB - ?E- ! " "" " L/ ! + "- + &% ! "!& (*"

49 Article 1

B ) + ! N " 8" + : ( -8" / ) ! "" - B + ( +'/ BBB+ )! "- / = " '+ "8+- / + " +C BB ! ! " + +$ " (*"# BBD) -'+ BE- - -! "/& (*" BBG) " " -+ 8 "& / ." ! "! + ( ) ! " +. ?( B ) "/"+ /(/

- + ! C B) ! "! / 93A (3 JJD) " ( 7 #N BB)* '? B( "#N BBJ)= " +- "-"' + * ! BBB ""- =8""G M - G + :"" -W?W"

+ '"-! -" + + + - +" ! + + ""-- + - / (* B BK )K M/ +-"'" + --++ - ! "! "+ -L'" -' / + 9( ) . 8" S$ "(.S$) "()/+!: + / ( $) + "+ ' - + "-/" +! ' " " (+ "+/ 'K / B K " B ) /"" "--- (+--' N + K BB>K / B K3L ! #& + B ) ! "+ ! " ' - ( B ) !X.S$ " " '" L'! / "+ (3L ! #& + B ) "! "'" "-/ ' /:!+ " + : ( BBG) " : ( " ) ! -

50 RESULTATS

" - ' " M/ :"- " + " + + - ( BBG).S$M " M '"(3L ! # & + B ) !"" '- /& .S$ ' ' + "+/- ( " B ) !8"-- - ' "- + 8 "&% +& / "& " "! =+ " + 8D( : BBG) " '= '" + <">(& BBD) / -8+ BB & / !"'" +& / (M B ) !!"" -?B+ !- :' (" ") +'/ BBB+ ! : : ! /-8" "! " + " - N=:"- '+ " " ' ! "($ B ) " "-/" ! " - : + ' B> ( " BB) & / -& ! " " ' !"X /!: / / +'" - "L'"""'" ' / +- (< #" / BBD);!: "" + J?E / / !*; M ! "/+ / + " "-- ( JJ) + / " + (D>EGG8 < #" / BBD)= " /!: $ ($ BBB) +" ( B"'" ) /+J?E "JJE " ++ + "

+ A , =' + "!*; "' / " " ++ /! -!+ + (B B)K "+/ - ! ' M N *' / "/ "+ " ! " +"= +" '/ BB: " " +" " ! " " - ? "'"

51 Article 1

+ BBB = " "' / " "' / (< )

/ (<) " /"8()(& " B )($ B ) + '""/; +<++"' /"! / - ! + / "- - " " + - + - "-- + ( BB>) =8"-- -"-- "' /" ' /++"" ?B?B: " -"'" / B = "++ - + + " % &/* =''(5 BB?).+ " + ! " +.+ * 8. ' ( 9YY"' /- +YZ Y+"+) "+" ! ! " . = -" ! JJJ +- - ! + " " + -"++ " =8"" + & "&- !+! - ([?) + ?!"- / ( ) " + + ! """/

% ! :" ! + !8"! "'+ 3 ! " - " +- " " + ( # B ) " ! " "- *; + / " "- / (*/ "" K&"+ /& # " B ) ! - /"'+ ! "'+ - - !! " ! " " " ! BBJ?\ BBB "BB JJ\BBB '/* + / !""& ' D (*" B ) + "+ 8":(*" BBG) " S7 '- +! "" / " ! " !"" - B + + '/ BBB + - / = "'+ "8+- / + " +CBB + !" + +$ " -'+ BE

52 RESULTATS

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

53 Article 1

+ ("< ".) " (" ) " "--( +" ) " ! "B? +:++K() "-- ! """ " " " = " "-- - " ' "-- - ( 9YY!!!!"+Y"! + -" ) " + + /"! " " /- ' (

54 RESULTATS

$X; !+ " + 8' ( BBG) ! "-8 !L+ ( B 7! B ) 8N '/!! /" X-" " "- :+" "/ X"-"' ( B B)= ""- -" BB " " + " '" =:" - BB " "/ BBB "'"++" ' B +" ! " " - B - 7.$ + " !'" /+" " B + + ( BB>), - 7.$ (3" B ) '" " - " - ! :!"+ "+ / " "" +"""

( $' B ) = -+" / / " " +- ' + () / " "+ - 8 "-/ + ! " ($ B )! /;& '(9YY" "Y >B>Y B ) /7" /(7) (7! B ) "-" " 8 ' ! '"- / '"+ "-:! " - % X ' "'!-" ! "!- " "9 --" ! "" " --" +' '7 $ ( 7$) ( BB>) 7$ '" /- " ! " 7$ / M !++- " " " -" " /"' " ! + O & "'"/;& '(9YY" "Y >B>Y BB?J) ! 8" -&_ E( BB>)=-" BBB / +?BE- ! ' X= "-" -+ " / - . & + !

55 Article 1

+ !+ "+ " + /+ / " "( >BB: )( :# / BB?) ! "(. "< " ) "+ :/ " ( N" : BBK"6+N4N# / BB>)= :!"+ !' / " - -+ " - " : +" - -- ="-"-+ +! 9 ( )M" 7.$ + - /+-"-- " (>-> 7.$ + GY> Y> " Y>) "-"-- / - . "< " " / - ! : - -+ ! "-"-- -+ (3+ ) " () "-+ - "'" 7.$(3+&)K!" - ! : (. "< " " /)- 7.$""/3"'" 7.$ --+ ! " " - "-/ + (>-> 7.$ + GY> Y> " Y>)%-- !" 8" '/ " " "

> '-+ = "!" !$77 "% 7.$ -8 " 7.$ , +;"-/ '-+ " "'+ K !' " " 8 " + / /+- + - " 9 " "- (-+ B B)

+ ;! M ""+ "/!" ".& :! 05?GB?GB05?GB>G( "" - )K "" M !"! "" - .& : ( "" - ) % B M ! " + G "-- / 3" ++ " " " !

<0S. " - - " " & / -

56 RESULTATS

K! " " " /+ / 8 8 !/ . ."! ' /+ ! /" !&% " ( ' & ?J?)% ! ! " 9 "& (3+)$ " "" 8! " + ( ! C>>E " CBDG)3 + ! "8 '/-"'" -(" + ) " ! " " + GK " "- ! " " " - ? G ? "" "'" - ! % " G """'" - " $ " & "? + (& &?) ! "( CJDE B)&? " + + - "& " + & ! ! " " ( JJE B) +&& ! ! " ( ?E BG)& " & "& ( DJE BJ>)&! "8 '/- % "'" K + + """'" -"-- " + ." +"- ?D>E!' + ( "" - ) ! L " ( "&)! "/ " -DBE( "" -); / - + " 3A -

"+ 8 ! + ' " +- 8 +& ! ! " +- -! + !+ N ( G "&?) - ! ++ "+ 8 - + 8- +&

= "" M"-.S$ / (3+ )% :" -S"! +- / / (& (_8 BG 8 " <">(_D8 BD " & / (_ >8 BG '/)$ "-"(.S$+ "- +"'" )! /+ " K D

57 Article 1

!"" + "-- ! " - "+ + 9 !'" ( + ) + "!& (-"' 9 ) ! +! / "" (-" ' 9 ! 8 B !& / ) *-- "+ "+ + &? " G !.S$ / + & - + ! (3+)

9 /-""' / " /

+ < <* J?E$% J?E$% G G >B> B BJ> BBBB? G> G>J J BB >? BB GBBDJ & G >B ? BGB BBBB?D >J?JDBB BBDB; BB?JDB? & B ?G? B BBBB? BJ; B?J> ?D B; B DDD>B & ??? ? B > BBBB B> B>>?GB?> BBD> ; BBGGB?DD &? G BD B?D BBB >D J>> J>> DG? BB ? BB??GBJD ' 9Z[BB?KZ[BB K;Z +- KZ- M K<Z- / K<"Z< /

"' /KZ""' /K3")3W 3 KZ 7 #N K$%Z$-"'

$ ' """+(3+)9 J?E "-!""!: " JJE%- 8- " ! -+D""!: K 8- " ! ? ! ""!: + " .S$ = -" ! /!: 9( ) ' + "-"'" " -! / -! / "' / () + / "'" - " & ! / " / -+ :!:(3+)

m mm k m k k m mm m m n m km m k m k kk m m km m mm m m m n mk m kk o o ! k " m o "! m o k "" #!# m $#$ k %% m

58 RESULTATS

GMYC TCS

MrBayesRAxMLBEAST95 99

LX1485 (MA) A1 LX1481 (MA) A2 LX1496 (MA) A3 LX1498 (MA) A4 GQ279225 (MA) LX1140 (IP) A5 LX1477 (MA) LX1286 (IP) LX1323 (IP) LX1603 (GR) LX1594 (IP) LX1268 (IP) LX1272 (IP) LX1317 (IP) LX2212 (GR) LX1143 (IP) A6 LX1561 (MA) LX1352 (IP) LX2264 (IP) LX2225 (GR) LX2342 (IP) LX1326 (IP) LX1542 (MA) LX2391 (IT) LX1336 (IP) LX1351 (IP) LX2211 (GR) LX1601 (GR) B1 LX1472 (MA) LX1712 (GR) LX1342 (IP) LX1941 (IP) LX1618 (IP) B2 LX1242 (IP) LX1264 (IP) LX1683 (IP) LX1146 (IP) B3 LX1148 (IP) LX1473 (MA) LX2267 (IP) LX1175 (TR) B4 LX1659 (BI) LX1048 (TN)

LX1209 (BI) LX1745 (CR) LX1753 (CR) LX1199 (SA) LX1938 (IP) LX1752 (CR) LX1219 (BI) LX2216 (GR) LX1738 (CR) LX1029 (TN) B5 LX2235 (GR) LX1273 (IP) LX2447 (SC) LX2059 (TN) LX2073 (TN)

Mr. Bayes BEAST LX1933 (TR) > 0.9 > 0.9 LX1592 (IP) 0.02 RAxML LX1177 (TR) > 75 LX1608 (LE) LX1759 (CR)

59 Article 1

+ A , / ! "" " " ! + ! !" " ( G & & "&?K " "" -) /

+ :" ++ - "' /(< ) " /

(<)K!' /8"+' - /"8() / -"'" + !" ++ - +& "& !+' - "' / ! "% " & % " '/ + G " &? !" ++ ! + ' ' " + *-- N - "" ( !)K" + -"'" (""/ ; + ) + ' 8 ! + & ! + ' - N"+ ". !"! : ! + "++ " - + G & & "&?([B) ! +- 8- +&?( )$' / ! + ? " ! " + " +- (_B? [BB?K ) ++ + / ++ / "

9 - / -"! BBBB +- - !+" "+ -"++ " + ? B?? BB G G BB?> B DG & BBB B> & B ?J BB?J &? B [BBBB

! mm m ! n m m m km m m m k m& mk m '(# km k ) 'm *+'m k ,+'m m m m- mm m km m m m k k (# m ) m ) m (#m m & m mp&../n m m *+'m k ,+'m m m

60 RESULTATS

Method 1 21 Ka min 1/8 CCSM CNRM COSMOS min 4/8 GISS IPSL MIROC min 6/8 MPI MRI

min 8/8

min 1/6 6 Ka CCSM CNRM min 4/6 IPSL MIROC MPI min 5/6 MRI

min 6/6

Present min 1/8 CCSM CNRM COSMOS min 4/8 GISS IPSL MIROC min 6/8 MPI MRI

min 8/8

Refugia min 1/8 min 6/8

min 4/8 min 8/8

Method 2 Present 6 Ka 21 Ka Intersection of 3 time slices Refugia min 1/8 CCSM CNRM

min 4/8 COSMOS GISS

min 6/8 IPSL MIROC

min 7/8 [8/8 not possible] MPI MRI

61 Article 1

% *'+ "--" ""+ " ( "" -?)& / !' " " L"' - ' " - + " !! L " ( %& 3+)! " " B?G (J?E<*9BB>D) + - " JG> ( GJ>)! $

) X" -"'" 7.$ !"'-"" - ! " / +- 7$ ( [BB?K "" -G)" ! 7.$ ( "" -D) ! "-" % % / . ! :/ - "' " " "-- !" + " -8 ! "-" :/"%7$ "% + ' / " " ! (++" / 7 / )! 7.$ '+" (+ " ) " + "& : K + ! "" / ( * X + "-" ! "-" "+"<( "" -D)7 ! ! !89 +%"-" " +! 7.$ 7 / " " - + ' /' "!.L-%7$ -+ " ! --"/' - !" !"" $ " ""<L . " -" ! " " (% % / & : ) " " +""8 8 '+ - !! +/ ""< !"-" "+ ." 7.$ + ' / "' " !" " ! - " - "/

62 RESULTATS

(a) A1 A1 (n=2) A2 (n=1) A3 (n=1) A2 A3 A4

A4 (n=6)

(b) A5 A5 (n=2) A6 (n=142) A6

(d) B2 B2 (n=36)

B3 (e) B3 (n=10) B4 (n=42) B4

(f) B5 B5 (n=61)

Figure 4 Distribution map and haplotype network for each lineage. Colors on haplotype networks correspond to colored areas on the maps to the left. Haplotype networks are not to the same scale among lineages. Each circle represents one haplotype, and colors correspond to frequencies of origin region of the haplotype (n = number of individuals). Note the two contrasting phylogeographic patterns, with lineages A1 to A5 restricted to one or a few well-structured populations, whereas the lineages distributed across the Mediterranean Basin generally lack geographic structure, likely a consequence of human-mediated dispersal.

63 Article 1

. A A -"'" 7.$ ! !8-%& % " "--'+ "- 7.$ (-8 "" / - '+ ! 7.$ + - ) ! -+ " / ! "!%K ' -+ !+ !"+ "! "!%(3+?K!% !)*"+'- +/-"- 7.$ + L "" " "-+ !"-"(3+? )= + + " ! "(3+? ") " ! "-" -+ 8- " - " ".- -. =" ' - +/(3+? - +) -+ ! "! " / + " -% ($ " . $ "NY + + O +K I" # * " BBJ) & % " / !"-" -+ '" + (3+?" ")K -/!'" " (8 ) ,: !

" -! - --" !"-" -+ 38 / - (.)!'" / '-+ K " - "' !'" -+ /" + 7.$ +'

%. (.$)

' , ="+*; +"' / " & "' //* (B B) " + - " + -- - - "' / - " " ' " ' / + (3+) " / / +/-'"( *; ) '" ($ B ) "-" + " : -- " - 8 ++ ' "+/ "' : ( B ) " /++ '"8 ! ""

64 RESULTATS

" ;' ' ! - / 8 - "'+ " + " ' " ! " "- + "' /

! mn mm m k n m n m m& n m m- k 0 >

."' / " "-/ + + ! " - +/+ (3+ ")& "/ !-"! + /++ 9 +- ' + ! ! " ! + " " " " & + / :++ 3 + ( ) " " +! - (3+) "+"'" - -" Q "R/*

65 Article 1

(B B) + + ' 8/(!?BBB:) /8 :++"'+(CDE "" -) + ! + /"8' / "+ + ( "" -) -"+"'+ " /"-- +

8/ /8 "/+ - + "! " /" " +! ? + / " +"' / '/ "!" ! + ! " ! " (+/+ B ) "!

-$ /% "( # B ) N+ "

"+- O /(K "" ) "'" !' + + !" " " (+ G " & + 3+)< /!: - + ' : ! / " + "'" " " " & "-! / ""'" - -! 3 + "8 ! / " ++ / ! ! : ! + " ++ " 38 /- G -" /- " & (3+) + "Q% "R/* (B B) ! " " "'" - + / /- &++ ! "& 8 8 +& ! "'" -"% % + & & "&? "'" ! / !" " " G + + : + "' / ! ++ / ( "" - " ) +& 88 - 8 " -" ! ' +' 8 ' +!"" !-""'" - + /? "/CBBB:( + - ++"/ : -& " + ) * + ! " " " " "-- " (< B "- )$+ -

66 RESULTATS

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

67 Article 1

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

68 RESULTATS

B ) "++ N" '(3+?) + ? " & ' " " "% ! /- -+ / + " % " <!' ? + !"/" " " . "' / " " + + - + + -+ " ++"' /( ="# 8 BB ) - +&( "" -) !+ + "' / -"& % " "-" / -+ (3+ ")<!' + G & & "&? !" " " "" ! / "!+"' / " + '-+ K + / '+ "- +"" -+ (+/ % ) M (+ " " ) ' " ++ + 8 ' + " " / ' M

$) -(.$)

% "/!"" ' / + ! " & " *; " . "' / ! " " -/ "!-"! +/++ 9( ) + -" ' + !! " ()! + " " "" & + / :++ &/ + - > 7.$ ! ! "-- -+" "+ ! " --+ !" / - ; ! --'"++ '! - ' -+ !- " ""+'- +/ "- + +" +-+ " ++ /+; !/++ !-" -/ - ! -+- +"' / /+ " / "! "/ '+ "+ " " ' "" - !" &

69 Article 1

!)$#-%')

= : ! "+/ " ! -"!: '"" - "/ / " &"+ / + . ' +' % 2N 0&+ : S : * 6N 5 +" O7 ' O " ";S= : " " - +" - "/ = + -;& ' " -'"+ 3"+- ! '""/$.BB>B>?L( "$ / +6 ) ! "/ 3%+ - "$ / +6 (& BBJB ?>D ) " ! " / ,' / - 0 - . " 3! " ;3.0 3! = :!"+="$ + A =:+.$""+ ! -$%? " ! : "++ ( "+%- "" -- )-"+ " :+" '

- *B . 9 < < /-" 78-",' / ;!S: +; 33 "$ &: ' N; * '"5 . " +B B7+ " 8 - " + ( !" '-! ,$ >D9 DB >B & '; & 'O 5INO '" ;+ 25 O 3B - + "+ " " "+& 9 > B > J & ". 0 " & ; ' I 34" N & :: +; & $-* &&N"$ -- O O + ' OB 8 /-'9- P /"' - -" + /" ' $ " ->B9B > &-".5 $ &" / ; N& $ * BB> /+ - " " " ! ! ."! ' $ &% J9?>?? &" 5 5 5S&" ".&-B B " +9&+ *' / " ""78-",' / ;!S: &+ JGJ; +/"" "A% ( ) G9 GG & " $ 5 M* "M' &0BBD +"'+ +/+ " - ?G9D D? $ N ";BBG / ++ / "'-. +: ( 9.::" )" " +" *; M $

70 RESULTATS

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

71 Article 1

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

72 RESULTATS

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

73 Article 1

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86 (Last GlacialMaximum,mid-Holocene,andpresent). slice time each in model circulationgeneral each forresults modelling niche Ecological MRI MPI MIROC IPSL GISS COSMOS CNRM CCSM Present Ecologicalnichemodellingresults for eachgeneral circulation model. mid-Holocene (~6Ka) Last Glacial Maximum (~21Ka) Last GlacialMaximum RESULTATS 87

CAPÍTOL 2

Diversitat del gènere Loxosceles a les Illes Canàries

Article 2 Planas E, Ribera C. 2014. Uncovering overlooked island diversity: colonization and diversiﬁcation of the medically important spider genus Loxosceles (Arachnida: Sicariidae) on the Canary Islands.

Article 3 Planas E, Ribera C. Description of six new species of Loxosceles (Araneae: Sicariidae) endemic to the Canary Islands, and the utility of DNA barcoding for their fast and accurate identification.

Article 4 Planas E, Bernaus L, Ribera C. Development of novel microsatellite markers for the spider genus Loxosceles (Sicariidae) using next-generation sequencing.

Article 5 Planas E, Bernaus L, Sánchez-Gracia A, Ribera C. Genetic diversity is affected by Pleistocenic sea-level changes and volcanic activity in a spider of the genus Loxosceles endemic to the eastern Canary Islands.

RESULTATS

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RESULTATS Journal of Biogeography (J. Biogeogr.) (2014) 41, 1255–1266

ORIGINAL Uncovering overlooked island diversity: ARTICLE colonization and diversiﬁcation of the medically important spider genus Loxosceles (Arachnida: Sicariidae) on the Canary Islands Enric Planas and Carles Ribera*

Institut de Recerca de la Biodiversitat (IRBio) ABSTRACT and Departament de Biologia Animal, Aim Our aim was to assess the evolutionary history of the spider genus Loxos- Facultat de Biologia, Universitat de celes on the Canary Islands. We unravelled its present diversity within the Barcelona, 08028 Barcelona, Spain archipelago, and investigated its origin, mode and tempo of colonization to and between the islands using a phylogenetic framework. Location Canary Islands, Madeira, Iberian Peninsula, North Africa, Mediterra- nean region, Guinea. Methods We conducted extensive sampling across the Canary Islands, and examined the phylogenetic relationships among the Canary Island representatives of the genus Loxosceles and with regard to species from western Africa and the Mediterranean Basin. We used an evolutionary criterion (general mixed Yule coalescent) to delimit the evolutionary lineages, and applied fossil and biogeographical calibration points to estimate dates for major cladogenetic events within the Canary Islands using a Bayesian framework. Results Phylogenetic analyses revealed the existence of a well-supported clade formed exclusively by Canarian Loxosceles specimens, comprising seven allopatrically distributed evolutionary lineages. Major dispersal events between the islands occurred during the late Miocene. Representatives of the cosmopolitan Loxosceles rufescens were also found on the archipelago.

Main conclusions We have revealed the existence of an overlooked endemic group of medically important spiders. The pattern of diversity of this group ﬁts well with the general dynamic theory of oceanic island biogeography, where maximum diversity is found on islands of intermediate age. The colonization pathway of the group is compatible with a stepping-stone model. Between- islands dispersal was the major driving force for diversiﬁcation in the group, but a few within-island speciation events were also inferred, such as on Gran Canaria, where the Roque Nublo volcanic event acted as a vicariant agent, promoting the split between the two Gran Canarian lineages. The recently intro- *Correspondence: Carles Ribera, Institut de Recerca de la Biodiversitat (IRBio) and duced L. rufescens is cohabiting with the endemic lineages. Departament de Biologia Animal, Facultat de Keywords Biologia, Universitat de Barcelona, 08028, Loxosceles Barcelona, Spain. Biogeography, introduced species, island endemics, island evolution, , E-mail: [email protected] Macaronesia, Mediterranean Basin, phylogeography, Roque Nublo.

of this diversity, however, is a necessary step in conducting INTRODUCTION studies of such islands, and understanding the origin of an Oceanic islands are excellent natural laboratories for reveal- island group of taxa is of primary interest (Emerson, 2002). ing the evolutionary mechanisms involved in generating As oceanic islands, the Canary Islands have never been in diversity (Emerson, 2002; Gillespie & Roderick, 2002; Whit- contact with a continental landmass; consequently, all the taker & Fernandez-Palacios, 2007). Reliable documentation terrestrial biota encountered there can be traced back to an

ª 2014 John Wiley & Sons Ltd http://wileyonlinelibrary.com/journal/jbi 1255 doi:10.1111/jbi.12321 93 Article 2 E. Planas and C. Ribera original dispersal event, either active or passive. Two princi- 2004, and Swanson & Vetter, 2006, for reviews), mostly pal areas have acted as a source for the Canary Island biota: caused by the action of the enzyme sphingomyelinase D the neighbouring North Africa landmass and the Iberian (Binford et al., 2009), which causes the clinical condition of Peninsula (Juan et al., 2000). However, in some cases, more loxoscelism. closely related groups are found on other islands of the Mac- In the Canary Islands, the only representative of the genus aronesian archipelago, i.e. the Azores, Madeira, Selvagens presently recorded is L. rufescens, which is considered as and Cape Verde, or even in more distant regions, such as ‘probably introduced’ in the Canarian species checklist the New World (Carine, 2005). (Macıas, 2010). Loxosceles rufescens is considered to be Diversification patterns and colonization pathways are also cosmopolitan because it has probably been transported a central subject in the study of the evolutionary history of worldwide by humans, but its native distribution appears to island taxa. The use of neutral markers for phylogenetic be the Mediterranean region (Gertsch, 1967; Duncan et al., inference has greatly improved our present understanding of 2009). However, Duncan et al. (2009) included two distinct these island biogeographical patterns (Juan et al., 2000; Loxosceles representatives from Gran Canaria in their analy- Emerson, 2002). In addition, molecular data provide infor- ses: one that was molecularly indistinguishable from the mation on the time frame of the diversification of island Iberian L. rufescens, and another that was morphologically taxa. Oceanic islands have been widely used as calibration and molecularly divergent not only from L. rufescens but also points for divergence time estimation, given that the age of from any other Loxosceles species. subaerial volcanism that formed the original island provides In the present study, our aim was to (1) investigate the a maximum time limit for the evolutionary history of its diversity of the genus Loxosceles in the Canary Islands archi- biota (but see Heads, 2011). pelago, and (2) infer a phylogenetic framework for investi- Volcanism also plays a key role in shaping within-island gating the biogeographical patterns and evolutionary history diversification. Several studies have shown that phylogeo- of Loxosceles. We conducted a thorough sampling of Loxosce- graphical patterns in organisms such as spiders, beetles and les in the Canary Islands, and obtained DNA sequence data lizards closely match well-dated pulses of volcanic activity. In from those specimens plus representatives from related geo- the Canary Islands, these effects are most evident for recent graphical regions, to infer a phylogeny and to delimit distinct volcanism events (e.g. Emerson et al., 2006; Bidegaray-Batista evolutionary lineages. Molecular data were used further to et al., 2007) but effects of older volcanic activity (e.g. the propose a time frame for the diversification of the group, eruption of Roque Nublo on the island of Gran Canaria) can using fossil and biogeographical calibration points. Because still be found (e.g. Contreras-Dıaz et al., 2003). of the medical importance of Loxosceles spiders, the results of Spiders are among the most diverse of the arthropod this study could have far-reaching implications for the orders, and have been shown to be a good model for the islands’ epidemiology and treatment of spider bites. study of island biogeography (Hormiga et al., 2003; Gillespie, 2004; De Busschere et al., 2010). The present-day knowledge MATERIALS AND METHODS of spider diversity on the Canary Islands is reasonably com- plete (Cardoso et al., 2010), including an updated checklist Sampling and molecular data collection (Macıas, 2010) and several taxonomic revisions and phylogeographical studies (Wunderlich, 1994; Arnedo et al., 2001, Loxosceles specimens were collected between 2009 and 2012 2007; Platnick et al., 2001; Lopez-Mercader, 2005; Bidegaray- and complemented with material provided by colleagues. To Batista et al., 2007; Dimitrov & Ribera, 2007; Dimitrov et al., assess their relationships with continental representatives of 2008; Macıas-Hernandez et al., 2010, 2013a, 2013b). The the genus, we included known species present in the Medi- information gathered for these studies, along with research terranean Basin, namely two individuals of Loxosceles mrazig conducted on other terrestrial invertebrates, has led some Ribera & Planas, 2009 from Tunisia (Ribera & Planas, 2009), authors to propose emerging evolutionary and biogeographi- and three of L. rufescens from the Iberian Peninsula, illustrat- cal patterns (Juan et al., 2000; Cardoso et al., 2010; Cameron ing the main evolutionary lineages found across the entire et al., 2013), generating greater interest in such research. Mediterranean (Duncan et al., 2009; E. Planas et al., unpub- The spider genus Loxosceles Heineken and Lowe, 1832 is lished data). In addition, we included two individuals one of the two genera that form the family Sicariidae Keyser- belonging to Loxosceles foutadjalloni Millot, 1941 from Gui- ling, 1880. Loxosceles consists of 107 species (Platnick, 2014), nea (Duncan et al., 2009), two L. rufescens from Madeira mostly distributed across the Americas and Africa (Gertsch, and three Loxosceles aff. rufescens individuals from the Anti- 1967; Gertsch & Ennik, 1983; Binford et al., 2008). Most of Atlas region of Morocco (E.P. & C.R., unpublished data). the species show a narrow, circumscribed, distribution, Phylogenetic trees were rooted with one representative of although a few exceptions exist, for example Loxosceles laeta Loxosceles vonwredei Newlands, 1980 from Namibia (Binford (Nicolet, 1849), Loxosceles reclusa Gertsch & Mulaik, 1940 et al., 2008). and Loxosceles rufescens (Dufour, 1820). Loxosceles spiders are For all the individuals, we amplified a partial fragment of well known for their medical relevance; their bites can cause the cytochrome c oxidase subunit I (COI). Additionally, one severe dermonecrotic lesions in mammals (see da Silva et al., individual per locality was genotyped for three more gene

1256 Journal of Biogeography 41, 1255–1266 ª 2014 John Wiley & Sons Ltd 94 RESULTATS Loxosceles on the Canary Islands regions, namely one mitochondrial fragment spanning the 30 GUI 1.3 (Silvestro & Michalak, 2011) in rapid-hill climbing half of the large ribosomal (r) unit (16S rRNA), the transfer mode. A GTR+G+I nucleotide substitution model was (t)RNA leucine (L1) and the 50 end of NADH dehydrogenase applied to each of the partitions corresponding to the best (nad1), and two nuclear genes, histone 3 (H3) and the inter- partition scheme selected, and a nonparametric bootstrap nal transcribed spacer 2 (ITS2). Primers are listed in Table 1 support analysis of 1000 pseudoreplicates was conducted. BI and polymerase chain reactions (PCR) were conducted fol- analyses were conducted with MrBayes 3.2 (Ronquist et al., lowing Planas et al. (2013). Raw sequences were edited and 2012). The best-fitting model was applied to each partition, assembled with Geneious 4.6.5 (Drummond et al., 2009). and all parameters were unlinked across partitions. Two All new sequences obtained were deposited in GenBank (see independent runs were performed to check for convergence. Appendix S1 in Supporting Information). For each analysis, four Markov chains were run (one cold, three heated) for 2 million iterations, saving trees each 1000th generation. The first 25% of trees in each run were Alignment, partitioning analyses and evolutionary discarded as burn-in following visual inspection in Tracer model selection (Rambaut & Drummond, 2007), after ensuring that the Mar- We performed sequence alignments using the online version kov chains had reached a stationary (average standard devia- of mafft (Katoh et al., 2005). We applied the G-INS-i tion of the split frequencies of the two runs < 0.01), and a algorithm (Katoh et al., 2005) for protein-coding fragments majority-rule consensus tree was generated from the remain- and the Q-INS-i algorithm (Katoh & Toh, 2008) for the ing trees. ribosomal gene fragments. Both algorithms were applied We used the tree-based general mixed Yule coalescent using the default options. (GMYC) model (Pons et al., 2006) to define species bound- We explored 11 alternative partitioning schemes using the aries. This method has been applied successfully for DNA- ‘user-specific schemes’ option in PartitionFinder 1.0.1 based species delimitation in different organisms, including (Lanfear et al., 2012). Partitioning schemes ranged from con- spiders (e.g. Hendrixson et al., 2012; Planas et al., 2013). sidering all the gene fragments together as one partition, to a Although GMYC relies exclusively on one line of evidence, highly partitioned scheme with each gene fragment consid- i.e. mitochondrial (mt)DNA, for species delimitation, it has ered as an independent partition and the third codon posi- been found to be accurate and conservative in groups with tion separated from the first and second in coding regions small effective populations and high divergence times (see Appendix S2). We used the Bayesian information crite- between species (Fujisawa & Barraclough, 2013), as is the rion to select among the partition schemes and evolutionary case in this study. models (Lanfear et al., 2012). We generated an ultrametric tree with beast 1.7.4 (Drum- mond et al., 2012) based on the COI DNA sequences. We used a coalescent tree prior with a constant population size Phylogenetic analyses and delimitation (Monaghan et al., 2009) and an uncorrelated relaxed molec- of evolutionary lineages ular clock with a molecular rate set to 1, as we were only We conducted phylogenetic analyses of the concatenated interested in the relative branch lengths. We performed two matrix using maximum likelihood (ML) and Bayesian infer- independent runs of 10 million generations each. The results ence (BI). ML analyses were conducted using RAxML 7.4.2 of the two analyses were inspected in Tracer to assess the (Stamatakis, 2006) through the graphical front-end RAxML- correct mixing of the chains (effective sample size,

Table 1 The primers used for the study of the spider genus Loxosceles (Arachnida: Sicariidae) from the Canary Islands.

Gene fragment* Primer PO† Sequence (50–30) Reference

COI LINF F GGGCNTGRTCWGGNATRATAGG This study CANF F GCDGGDGCTTCTTCDATTATRGG This study LJEF F TCARCATYTRTTTTGRTTTTTTGG This study GAYAR R GAAAATGHGCHACHACRTAATAAGTRTC This study CANR R GCNCCYATAATHGAAGAAGC This study LJER R TGTTGAAAYAAAATHGGRTCHCC This study 16S+L1+nad1 Lox16SF F CGCCCTGTTTAACAAAAACATCAC This study 16SR R CCTTTAACGAATTTGAATATA Hedin & Maddison (2001) H3 H3aF F ATGGCTCGTACCAAGCAGACVGC Colgan et al. (1998) H3aR R ATATCCTTRGGCATRATRGTGAC Colgan et al. (1998) ITS2 CAS28sB1d R TTCTTTTCCTCCSCTTAYTRATATGCTTAA Ji et al. (2003) 5.8SF F CACGGGTCGATGAAGAACGC This study

*COI, partial fragment of the cytochrome c oxidase subunit I; 16S, mitochondrial fragment spanning the 30 half of the large ribosomal unit; L1, the transfer (t)RNA leucine; nad1, the 50 end of NADH dehydrogenase; H3, histone 3; ITS2, internal transcribed spacer 2. †Primer orientation: F, forward; R, reverse.

Journal of Biogeography 41, 1255–1266 1257 ª 2014 John Wiley & Sons Ltd 95 Article 2 E. Planas and C. Ribera

ESS > 200). We then conducted a 10% burn-in of each ages (e.g. 20 Ma; Binford et al., 2008). To account for age analysis and tree combination in LogCombiner and uncertainty, we used an exponential distribution with a TreeAnnotator (Drummond & Rambaut, 2007). The ‘hard’ minimum bound at 16 Ma and 95% of the posterior GMYC delimitation was conducted with the R package density within 21.99 Ma (10% of the older age 20 Ma), fol- ‘SPLITS’ (Ezard et al., 2009). Both single-threshold (Pons lowing Hipsley et al. (2009). In oceanic islands, geological et al., 2006) and multiple-threshold (Monaghan et al., 2009) ages represent the time of the earliest possible colonization models were applied to the tree, and a likelihood ratio test and they can be incorporated as maximum bounds assuming (LRT) was applied to select the best model. that species formation was subsequent to island emergence. We calculated the genetic divergence of COI sequences However, further major assumptions are also implied in this [uncorrected pairwise (p)-distance] between and within the case, which when violated would lead to incorrect divergence different Canarian evolutionary lineages (i.e. GMYC clusters) dates and molecular rates (Emerson, 2007). We used uni- using mega 5 (Tamura et al., 2011), removing alignment form priors from 0 to the time of emergence of the oldest gaps in pairwise comparisons. available island for the internal Canary Island dispersal events: (1) age of Gran Canaria (14.5 Ma) for the most recent common ancestor (MRCA) of Canarian endemic lin- Molecular dating analyses eages; (2) age of the oldest emerged part of Tenerife We estimated absolute divergence times and substitution (11.6 Ma) for the subsequent node; (3) age of La Gomera rates using a Bayesian approach as implemented in beast (10.5 Ma) for the MRCA of Tenerife and La Gomera ende- 1.7.4 (Drummond et al., 2012), applying an uncorrelated mic lineages. Well-supported clades recovered in the ML and lognormal relaxed clock (Drummond et al., 2006). We com- BI analyses of the concatenated matrix and those well sup- bined one representative of each evolutionary lineage of our ported in Binford et al. (2008) were constrained in the beast ingroup (Canarian and Mediterranean) with individuals analyses. Two independent runs of 30 million generations belonging to the Loxosceles amazonica Gertsch, 1967; Loxosce- (sampling every 1000th generation) were performed for each les gaucho Gertsch, 1967; Loxosceles hirsuta Mello-Leitao, analysis. Convergence of the chains was assessed and burn-in 1931, L. laeta, L. reclusa and L. vonwredei species groups. We was conducted as above. estimated absolute divergence times by incorporating fossil and biogeographical calibration points. Three Loxosceles spe- RESULTS cies are described from Dominican amber (Wunderlich, 1988, 2004). We thus constrained the ancestral node of the Appendix S1 lists the locality information for all 134 Loxosce- L. reclusa group (including Caribbean and North American les individuals used in the study, 125 of which correspond to Loxosceles; Binford et al., 2008) to a minimum bound. The 46 localities in the Canary Islands (Fig. 1). age of the Dominican amber is estimated to be 16 Ma (Iturr- The concatenated matrix used in the phylogenetic analyses alde-Vinent, 2001), although some authors have used older consisted of 2269 bp. The total length of the mtDNA gene

Madeira Loxosceles rufescens

Loxosceles mrazig Loxosceles Canary Islands LZ11 LZ7 Lanzarote LZ6 LZ8 aff. rufescens LZ4 LZ9 Anti-Atlas La Palma LZ2 LZ5LZ10 FV10 LZ1LZ3 Tenerife FV7 FV9 TF3 TF1 FV6 FV8 GC10 GC9 FV5 GM4 GC11 GC8 FV1 FV4 Loxosceles GM1 FV3 GC7 GC6 GM2 GM3 TF2 GC5 FV2 foutadjalloni La Gomera GC4 GC2 Fuerteventura HI1 GC3 GC1 El Hierro Gran Canaria

500km

Figure 1 Sampling localities of Loxosceles specimens. Colours correspond to those used in Fig. 2. Locality numbers are given for the Canarian endemic clade and correspond to those used in Appendix S1. FV, Fuerteventura; LZ, Lanzarote; GC, Gran Canaria; TF, Tenerife; GM, La Gomera; HI, El Hierro.

1258 Journal of Biogeography 41, 1255–1266 ª 2014 John Wiley & Sons Ltd 96 RESULTATS Loxosceles on the Canary Islands fragments was 1600 bp (COI 942 bp, 16S 547 bp, L1 57 bp evolutionary model for each of these three partitions is pre- and nad1 54 bp) and of the nuclear (nu)DNA gene frag- sented in Appendix S2. ments 669 bp (H3 327 bp and ITS2 342 bp). The preferred partitioning scheme included three parti- Delimitation of evolutionary lineages tions: the mtDNA gene fragments, except the third codon and phylogenetic analyses position of the COI gene; the third codon position of the COI gene; and the concatenation of the two nuDNA gene In the GMYC analyses, both the single- and multiple-thresh- regions. Information on length, variable sites and the selected old models ﬁtted to the data signiﬁcantly better than the null

LX1051 (TN) Loxosceles mrazig 0.1 LX1054 (TN) L_fouta4 L_fouta3 Loxosceles foutadjalloni 0.85/59 LX1016 (MA) Loxosceles aff. LX1015 (MA) 1/96 LX1494 (MA) rufescens Anti-Atlas 1/100 1/100 LX1330 (PI) LX1376 (PI) 1/100 LX1351 (PI) LX1068 (GC) 0.56/- LX1593 (HI) Loxosceles LX1468 (MD) 1/97 LX1188 (LP) rufescens LX1469 (MD) LX1599 (GC) LX1180 (LP) LX1797 (TF) LX2292 (HI) LX1795 (GM) Loxosceles sp. 1/100 LX1774 (GM) 0.97/77 LX1776 (GM) La Gomera-El Hierro 1/97 LX1782 (GM) 1/100 LX1796 (TF) Loxosceles sp. Tenerife 1 1/98 LX1164 (TF) Loxosceles sp. Tenerife 2 1/91 LX1155 (TF) Loxosceles sp. Tenerife 3 LX1149 (GC) 1/100 LX1687 (GC) Loxosceles sp. LX1707 (GC) Gran Canaria 2 1/95 LX1159 (GC) 1/97 LX1704 (GC) LX1813 (GC) Loxosceles sp. LX1160 (GC) LX2306 (GC) Gran Canaria 1 LX1165 (GC) LX1703 (GC) LX1701 (GC) 0.98/78 LX1400 (FV) LX1392 (FV) LX1394 (FV) 0.87/45 LX1169 (FV) LX1391 (FV) LX1406 (FV) LX1422 (FV) LX1412 (FV) 1/95 LX1418 (FV) LX1420 (FV) Loxosceles sp. LX1457 (LZ) Fuerteventura-Lanzarote LX1425 (LZ) LX1464 (LZ) LX1427 (LZ) 0.98/90 LX1463 (LZ) LX1440 (LZ) LX1454 (LZ) LX1452 (LZ) LX1445 (LZ) LX1459 (LZ) LX1466 (LZ)

Figure 2 Bayesian inference (BI) tree inferred based on a concatenated matrix [COI, partial fragment of the cytochrome c oxidase subunit I; 16S, mitochondrial fragment spanning the 30 half of the large ribosomal unit; L1, the transfer (t)RNA leucine; nad1, the 50 end of NADH dehydrogenase; H3, histone 3; ITS2, internal transcribed spacer 2] of Canary Islands’ endemic Loxosceles and close relatives. Numbers next to nodes correspond to posterior probability values in the BI analysis and to bootstrap support in the maximum likelihood (ML) analysis. Next to each specimen code the geographical region is provided: MD, Madeira; TN, Tunisia; MA, Morocco; GC, Gran Canaria; TF, Tenerife; HI, El Hierro; GM, La Gomera; FV, Fuerteventura; LZ, Lanzarote; LP, La Palma; PI, Iberian Peninsula). Evolutionary lineages delimited with a general mixed Yule coalescent (GMYC) model are named and coloured. The tree was rooted with L. vonwredei (not shown).

Journal of Biogeography 41, 1255–1266 1259 ª 2014 John Wiley & Sons Ltd 97 Article 2 E. Planas and C. Ribera model (single, P = 0.0113; multiple, P = 0.0185) but they Within the L. rufescens s.l. clade (PP = 1, BS = 100), two did not differ significantly from each other (v2 = 0.7728, different clades were recovered in both analyses: one corre- d.f. = 3, P = 0.8560). Thus the simplest model (single) was sponded to specimens from the Anti-Atlas region (Loxosceles favoured. Seven clusters and seven entities were retrieved, aff. rufescens Anti-Atlas) and the other to the remaining from which four clusters and three entities were formed L. rufescens specimens sampled (L. rufescens s.s.). In the exclusively by Canarian individuals (Fig. 2). L. rufescens s.s. clade, the representatives from the Iberian The topologies of the analyses conducted with BI and ML Peninsula, specifically from the type locality of the species were almost identical. Differences between the two methods (Sagunt), formed a clade together with several individuals included the position of L. mrazig and L. foutadjalloni with from the Canary Islands (Gran Canaria, El Hierro, La Palma regard to the clade grouping the remaining representatives and Tenerife) and the two specimens from Madeira (PP = 1, (i.e. L. rufescens s.l. and the Canarian clade). However, these BS = 100), with a very low within-clade COI uncorrected relationships were not well supported in any of the analyses genetic distance (0.1%). The sister group of this clade (Fig. 2). The sister relationship between the Canarian clade included two additional specimens from the Iberian Penin- and the L. rufescens s.l. clade was well supported in both sula, and its relationship with the former clade was well sup- analyses [posterior probability (PP) = 0.97, ML bootstrap ported in both analyses (PP = 1, BS = 100). support BS) = 77]. The clade formed exclusively by Canarian representatives Molecular dating analyses was supported in both the BI and ML analyses (PP = 0.98, BS = 78; Fig. 2). Within the Canarian clade, all the repre- The dataset consisted of 37 terminals and 1107 bp from sentatives from Fuerteventura and Lanzarote clustered sequences of two mtDNA genes, COI and 16S, common to this together with strong support (PP = 0.9, BS = 96) and con- study and Binford et al. (2008). The estimated chronogram is stituted a single GMYC cluster. Similarly, the Gran Canarian shown in Fig. 3. The age of the oldest split within the Canary (GC) specimens formed a clade (PP = 1, BS = 95) that Islands’ endemic Loxosceles, which separated the Fuerteventura– included two GMYC clusters (GC1 and GC2), both well Lanzarote lineage from the rest of the Canary Islands group, supported (PP = 1, BS = 97, and PP = 1, BS = 100, for was estimated to be 8.4 Ma (95% highest posterior density, GC1 and GC2, respectively). These two evolutionary lin- HPD: 6.03–10.98 Ma). The split between the Gran Canaria eages from Gran Canaria showed a 12.5% uncorrected species and the westernmost lineages was estimated to be divergence for COI and were distributed allopatrically (see 7.1 Ma (HPD: 5.09–9.26 Ma), and the MRCA of the two spe- Appendix S3). In Tenerife, the three sampled representatives cies from Gran Canaria was dated as 4.42 Ma (HPD: 2.65– formed a monophyletic group (PP = 1, BS = 98) and were 6.41 Ma). The split between the Tenerife representatives and delimited as three independent GMYC entities, with the the lineage from La Gomera–El Hierro was estimated to be COI uncorrected genetic distances among them being 5.33 Ma (HPD: 3.73–7.14 Ma). The first split within Tenerife greater than 11.4% (Table 2). The representatives of La Go- was dated as 3.67 Ma (HPD: 2.45–5.12 Ma) and the second mera and the single specimen from El Hierro formed a split as 1.99 Ma (HPD: 1.1–2.96 Ma). The estimated mean clade (PP = 1, BS = 100) and a single GMYC cluster. Rela- mtDNA substitution rate (12.686% divergence per Myr) was tionships between the main Canarian lineages were all well comparable with other rates calculated specifically for spiders – supported. The Gomera–El Hierro clade and the Tenerife in oceanic islands (5.3–9.8% divergence Myr 1; Arnedo & clade formed a clade sister to the Gran Canaria clade, and Gillespie, 2006; Dimitrov et al., 2008; Macıas-Hernandez all of them were in turn sister to the Fuerteventura–Lanza- et al., 2010) and five times higher than the ‘standard’ 2.3 – rote clade. divergence Myr 1 (Brower, 1994).

Table 2 Mean uncorrected distances of the partial fragment of the cytochrome c oxidase subunit I (COI) between (below the diagonal) and within (diagonal, in italics) Loxosceles endemic lineages from the Canary Islands. The ﬁnal column and row shows the mean uncorrected distance between individuals from La Gomera (GM) and El Hierro (HI).

FV–LZ GC1 GC2 TF1 TF2 TF3 GM–HI HI

FV–LZ 0.044 GC1 0.152 0.028 GC2 0.148 0.125 0.046 TF1 0.153 0.134 0.146 – TF2 0.150 0.140 0.141 0.114 – TF3 0.172 0.145 0.154 0.126 0.123 0.001 GM–HI 0.148 0.130 0.132 0.106 0.120 0.136 0.017 GM 0.014

Abbreviations: FV–LZ, Loxosceles sp. Fuerteventura–Lanzarote; GC1, Loxosceles sp. Gran Canaria 1; GC2, Loxosceles sp. Gran Canaria 2; TF1; Loxosceles sp. Tenerife 1; TF2, Loxosceles sp. Tenerife 2; TF3, Loxosceles sp. Tenerife 3, GM-HI; Loxosceles sp. La Gomera–El Hierro.

1260 Journal of Biogeography 41, 1255–1266 ª 2014 John Wiley & Sons Ltd 98 RESULTATS Loxosceles on the Canary Islands

Macıas-Hernandez et al., 2013a). Similarly, this could be the DISCUSSION main explanation for the current relationships between the three putative Loxosceles species found in Tenerife. An overlooked Loxosceles lineage endemic Another putative species of Loxosceles was found on the to the Canary Islands islands of La Gomera and El Hierro. The individual from El The results of ML and BI phylogenetic analyses (Fig. 2) Hierro falls among the representatives of La Gomera, which revealed the presence of a new Loxosceles lineage endemic to in combination with the low genetic distances (1.4%) sug- the archipelago. The GMYC species delimitation method gests a recent colonization from La Gomera to El Hierro. El identified seven evolutionary lineages within the archipelago. Hierro is the youngest island in the archipelago (1 Ma) and All of the islands, except La Palma, harbour at least one line- its endemic fauna usually shows close affinity with that of La age. The distribution of the endemic Loxosceles diversity fits Gomera, e.g. Calathus spretus (Emerson et al., 1999), Pholcus well with the ‘general dynamic model of oceanic island bio- bimbache (Dimitrov et al., 2008) and Dysdera gomerensis geography’ (Whittaker et al., 2008, 2010), with maximum (Macıas-Hernandez et al., 2013a). diversity found on islands of intermediate age. This diversity hump-shaped pattern has also been found in other spider Origin and colonization of the Canary Islands genera (Cardoso et al., 2010). Fuerteventura and Lanzarote together with the two The most confident strategy for testing the monophyly of an neighbouring islets of Lobos and La Graciosa harbour a island group is to include all the closely related species from monophyletic lineage, recovered as an independent GMYC continental areas and neighbouring archipelagos in the cluster and separated by a 14% COI uncorrected genetic analyses (Emerson, 2002). We therefore included specimens distance (Table 2) from the rest of the endemic Canarian from Madeira, all the currently accepted Loxosceles species in lineages. Within this lineage, two monophyletic the Mediterranean (i.e. L. rufescens and L. mrazig) and L. groups were recognized, one distributed across Fuerteven- foutadjalloni from Guinea in our phylogenetic analyses. Spe- tura and Lobos and the other across Lanzarote and La cial effort was taken to include representatives of different Graciosa. lineages distributed across the Anti-Atlas region of Morocco The two putative Loxosceles species of Gran Canaria are (E.P. & C.R., unpublished data). This region has been distributed allopatrically. Their distribution ranges roughly shown to be a continental source for many groups on the match the two parts of the island that were less affected by Canary Islands, because of its proximity with the eastern- the Roque Nublo eruptive period (Appendix S3). A similar most islands (Juan et al., 2000, and references therein). distribution pattern has been found in other organisms, However, omission of lineages because of incomplete sam- including the gecko Tarentola boettgeri, the skink Chalcides pling or extinction of continental relatives is a common sexlineatus and the beetle genus Pimelia (Pestano & Brown, caveat, which should be kept in mind when studying island 1999; Contreras-Dıaz et al., 2003; Gubitz€ et al., 2005). It has biota. Nevertheless, the monophyly of the endemic Canarian been postulated that a total extinction of the biota occurred Loxosceles is clear from the results presented here, obtained in Gran Canaria as a result of the Roque Nublo volcanic from molecular analyses, and is also supported by the mor- event (see Anderson et al., 2009), although most of the cur- phological synapomorphies shared by all of the Canarian rent evidence obtained from molecular data and fossils does representatives (E.P. & C.R., unpublished data). This ende- not support such a mass extinction (Emerson, 2003; Ander- mic group is a sister to the clade formed by L. rufescens and son et al., 2009). There is no doubt, however, that this volca- some divergent lineages from the Anti-Atlas region, and thus nic event had severe consequences on the island biota, it could be hypothesized that the colonization occurred from especially for species associated with forest communities the north-western part of Africa between 11.3 and 8.4 Ma (Emerson, 2003). Divergence time estimates place this split (Fig. 3). However, for conclusive evidence, it would be (4.42 Ma) within the temporal framework of the volcanic essential to sample the region between the Anti-Atlas and pulse (5–3 Ma) (Anderson et al., 2009) and, therefore, it Guinea intensively, as overlooked lineages could still be appears reasonable to associate the Roque Nublo eruption found and shed further light on the biogeographical history with the vicariant origin of the two local Loxosceles species of of the group. Gran Canaria. In the Canary Islands, the stepping-stone model of coloni- Tenerife originated as three isolated massifs (Anaga, Teno zation has been postulated as the most common pattern of and Roque del Conde) that emerged as a result of indepen- colonization shown by several groups of organisms (Juan dent volcanic cycles. These three proto-islands were later et al., 2000; Sanmartın et al., 2008). The model predicts that joined as a consequence of the Las Canadas~ emergence, the colonization of an archipelago begins with a dispersal forming the current island (see Marrero & Francisco-Ortega, event from the continent to the geographically closest island. 2001, and Carracedo et al., 2002, for reviews). This complex Then a succession of dispersal events leads to the coloniza- volcanic history has shaped the diversity and distribution tion of the remaining islands. In this archipelago, where the patterns of several organisms, including the two well-studied islands closer to the mainland are also the older islands spider genera Dysdera and Pholcus (Dimitrov et al., 2008; (Fuerteventura and Lanzarote), this pattern also corresponds

Journal of Biogeography 41, 1255–1266 1261 ª 2014 John Wiley & Sons Ltd 99 Article 2 E. Planas and C. Ribera to the progression rule, which states that younger islands are duction). In our analyses, some of the Loxosceles individuals colonized from nearby, older ones (Funk & Wagner, 1995). collected in the Canaries were closely related to the L. rufes- Given the inferred relationships within Loxosceles representa- cens Iberian lineage from the type locality (Fig. 2). Addition- tives in the Canary Islands, we do not have any evidence for ally, the two individuals from Madeira fell within the same rejecting the stepping-stone model as the pattern followed by lineage. The genetic distance of COI within this group is these spiders for the colonization of the archipelago, in suc- 0.1%, which is much lower than the intraspeciﬁc divergences cession from the easternmost to the westernmost islands found in any other group. Altogether, this evidence points (Fig. 3). towards a recent introduction of L. rufescens to the archipelago. Most Canarian L. rufescens were collected in human-dis- turbed areas; they were never collected together with any of Loxosceles rufescens as an introduced species the Canarian endemic Loxosceles. La Palma is the only island Before the present study, the genus Loxosceles in the Canary where we failed to obtain endemic Loxosceles, and only Islands was represented exclusively by L. rufescens (see Intro- L. rufescens was collected, even from well-preserved habitats

Loxosceles vonwredei Loxosceles sp. (Wundergat) Loxosceles hirsuta Loxosceles spadicea Loxosceles laeta Loxosceles sp. (Bonaire) Loxosceles sp. (Pisco) Loxosceles caribbaea Loxosceles chinateca Loxosceles reclusa Loxosceles colima Loxosceles baja Loxosceles kaiba Loxosceles deserta Loxosceles arizonica Loxosceles apachea Loxosceles blanda Loxosceles variegata Loxosceles amazonica Loxosceles mrazig Loxosceles foutadjalloni Loxosceles aff. rufescens Anti-Atlas Loxosceles aff. rufescens Anti-Atlas Loxosceles aff. rufescens Anti-Atlas Loxosceles rufescens Loxosceles rufescens Loxosceles rufescens Loxosceles sp. Fuerteventura Loxosceles sp.Lanzarote La Palma Lanzarote Loxosceles sp. Gran Canaria 2 3 Tenerife 1 1 Loxosceles sp. Gran Canaria 1 Loxosceles sp. La Gomera-El Hierro 2 La Gomera Fuerteventura Loxosceles sp. Tenerife 3 2 Africa 3 El Hierro 2 Gran Canaria Loxosceles sp. Tenerife 2 Loxosceles sp. Tenerife 1 30 20 10 0 Ma

Figure 3 Chronogram of Loxosceles specimens obtained using beast calibrated with fossil and island ages (see Materials and Methods). The black dot indicates the node calibrated as a minimum bound with a Loxosceles fossil from Dominican amber. The Canarian endemic lineage is highlighted in grey. Bars at internodes correspond to the 95% highest posterior density (HPD). The inset ﬁgure shows the Canary Islands, with arrows representing dispersal events using the stepping-stone model of colonization. Numbers in the inset ﬁgure correspond to those used in the chronogram.

1262 Journal of Biogeography 41, 1255–1266 ª 2014 John Wiley & Sons Ltd 100 RESULTATS Loxosceles on the Canary Islands such as caves. Conversely, Fuerteventura, Lanzarote and La REFERENCES Gomera are the only islands where we did not collect L. ru- Anderson, C.L., Channing, A. & Zamuner, A.B. (2009) Life, fescens. death and fossilization on Gran Canaria: implications for In the Canary Islands, as in the other Macaronesian Macaronesian biogeography and molecular dating. Journal archipelagos and on oceanic islands in general, there are few, of Biogeography, 36, 2189–2201. if any, venomous animals. Therefore the introduction of Arnedo, M.A. & Gillespie, R.G. (2006) Species diversification L. rufescens to the Canary Islands is of medical relevance. patterns in the Polynesian jumping spider genus Havaika Although loxoscelism is rare, even in areas with high Proszynski, 2001 (Araneae, Salticidae). Molecular Phyloge- Loxosceles density (Vetter, 2003), the synathropic lifestyle of netics and Evolution, 41, 472–495. L. rufescens facilitates human contact, making envenomation Arnedo, M.A., Oromı, P. & Ribera, C. (2001) Radiation of by this species more likely than by the endemic ones. The the spider genus Dysdera (Araneae, Dysderidae) in the existence of two distinct evolutionary lineages of Loxosceles Canary Islands: cladistic assessment based on multiple data in the Canaries, the endemic lineage and L. rufescens, should sets. Cladistics, 17, 313–353. be taken into consideration in future health management Arnedo, M.A., Oromı, P., Murria, C., Macıas-Hernandez, N. policies. & Ribera, C. (2007) The dark side of an island radiation: systematics and evolution of troglobitic spiders of the CONCLUSIONS genus Dysdera Latreille (Araneae: Dysderidae) in the Can- ary Islands. Invertebrate Systematics, 21, 623–660. 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E.P. was Carracedo, J.C., Perez Torrado, F.J., Ancochea, E., Meco, J., supported by an FPI grant from Ministerio de Ciencia y Hernan, F., Cubas, C.R., Casillas, R., Rodrıguez-Badiola, E. Tecnologıa (BES-2009-015871).

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Molecular Emerson, B.C. (2003) Genes, geology and biodiversity: faunal Phylogenetics and Evolution, 66, 327–340. and floral diversity on the island of Gran Canaria. Animal Hipsley, C.A., Himmelmann, L., Metzler, D. & Muller,€ J. Biodiversity and Conservation, 1,9–20. (2009) Integration of Bayesian molecular clock methods Emerson, B.C. (2007) Alarm bells for the molecular clock? and fossil-based soft bounds reveals early Cenozoic origin No support for Ho et al’.s model of time-dependent of African lacertid lizards. BMC Evolutionary Biology, 9, molecular rate estimates. Systematic Biology, 56, 337–345. 151. Emerson, B.C., Oromı, P. & Hewitt, G.M. (1999) MtDNA Hormiga, G., Arnedo, M.A. & Gillespie, R.G. (2003) Specia- phylogeography and recent intra-island diversification tion on a conveyor belt: sequential colonization of the

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Hawaiian islands by Orsonwelles spiders (Araneae, Liny- Monaghan, M.T., Wild, R., Elliot, M., Fujisawa, T., Balke, M., phiidae). Systematic Biology, 52,70–88. Inward, D.J.G., Lees, D.C., Ranaivosolo, R., Eggleton, P., Iturralde-Vinent, M.A. (2001) Geology of the amber-bearing Barraclough, T.G. & Vogler, A.P. (2009) Accelerated species deposits of the Greater Antilles. Caribbean Journal of Sci- inventory on Madagascar using coalescent-based models of ence, 37, 141–167. species delineation. Systematic Biology, 58, 298–311. Ji, Y.J., Zhang, D.X. & He, L.J. (2003) Evolutionary conserva- Pestano, J. & Brown, R.P. (1999) Geographical structuring of tion and versatility of a new set of primers for amplifying mitochondrial DNA in Chalcies sexlineatus within the the ribosomal internal transcribed spacer regions in insects island of Gran Canaria. Proceedings of the Royal Society B: and other invertebrates. Molecular Ecology Notes, 3, 581– Biological Sciences, 266, 805–812. 585. 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(2008) Infer- woodlouse hunter spiders (Araneae, Dysderidae) endemic ring dispersal: a Bayesian approach to phylogeny-based to the Macaronesian archipelagos. Systematics and Biodi- island biogeography, with special reference to the Canary versity, 8, 531–553. Islands. Journal of Biogeography, 35, 428–449. Macıas-Hernandez, N., Bidegaray-Batista, L., Emerson, B.C., da Silva, P.H., da Silveira, R.B., Appel, M.H., Mangili, O.C., Oromi, P. & Arnedo, M. (2013a) The imprint of geologic Gremski, W. & Veiga, S.S. (2004) Brown spiders and loxo- history on within-island diversification of woodlouse-hun- scelism. Toxicon, 44, 693–709. ter spiders (Araneae, Dysderidae) in the Canary Islands. Silvestro, D. & Michalak, I. (2011) raxmlGUI: a graphical Journal of Heredity, 104, 341–356. front-end for RAxML. Organisms Diversity & Evolution, Macıas-Hernandez, N., Bidegaray-Batista, L., Oromı, P. & 12, 335–337. Arnedo, M.A. (2013b) The odd couple: contrasting phy- Stamatakis, A. (2006) RAxML-VI-HPC: maximum likeli- logeographic patterns in two sympatric sibling species of hood-based phylogenetic analyses with thousands of taxa woodlouse-hunter spiders in the Canary Islands. Journal of and mixed models. Bioinformatics, 22, 2688–2690. Zoological Systematics and Evolutionary Research, 51,29– Swanson, D.L. & Vetter, R.S. (2006) Loxoscelism. Clinics in 37. Dermatology, 24, 213–221. Marrero, A. & Francisco-Ortega, J. (2001) Evolucion en islas: Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. la metafora especio-tiempo-forma. Naturaleza de las Islas & Kumar, S. (2011) MEGA5: molecular evolutionary Canarias: ecologıa y conservacion (ed. by J.M. Fernandez- genetics analysis using maximum likelihood, evolutionary Palacios and J.L. Martın Esquivel), pp. 133–140. Turquesa, distance, and maximum parsimony methods. Molecular Santa Cruz de Tenerife, Tenerife. Biology and Evolution, 28, 2731–2739.

Journal of Biogeography 41, 1255–1266 1265 ª 2014 John Wiley & Sons Ltd 103 Article 2 E. Planas and C. Ribera

Vetter, R. (2003) Diagnoses of brown recluse spider bites (lo- fossil and extant taxa. Beitr€age zur Araneologie, 3A,1– xoscelism) greatly outnumber actual veriﬁcations of the 1908. spider in four western American states. Toxicon, 42, 413– 418. SUPPORTING INFORMATION Whittaker, R.J. & Fernandez-Palacios, J.M. (2007) Island biogeography: ecology, evolution, and conservation, 2nd edn. Additional Supporting Information may be found in the Oxford University Press, Oxford. online version of this article: Whittaker, R.J., Triantis, K.A. & Ladle, R.J. (2008) A general Appendix S1 Study material. dynamic theory of oceanic island biogeography. Journal of Appendix S2 Partition schemes. Biogeography, 35, 977–994. Appendix S3 Distribution of Loxosceles species from Gran Whittaker, R.J., Triantis, K.A. & Ladle, R.J. (2010) A general Canaria. dynamic theory of oceanic island biogeography: extending the MacArthur–Wilson theory to accommodate the rise BIOSKETCHES and fall of volcanic islands. The theory of island biogeogra- – phy revisited (ed. by J.B. Losos and R.E. Ricklefs), pp. 88 Enric Planas is a PhD student at the University of Barce- 115. Princeton University Press, Princeton, NJ. lona, studying the biogeography and systematics of Loxosceles Wunderlich, J. (1988) Die fossilen spinnen im Dominikanis- spiders in the Mediterranean and the Canary Islands. chen bernstein. Beitr€age zur Araneologie, 2,1–378. Wunderlich, J. (1994) Zu okologie, biogeographie, evolution Carles Ribera is an associate professor at the University of und taxonomie einiger spinnen der makaronesischen in- Barcelona. His research focuses on the taxonomy and bioge- seln (Arachnida: Araneae). Beitr€age zur Araneologie, 4,1– ography of spiders of the Mediterranean Basin and oceanic 778. islands. Wunderlich, J. (2004) Fossil spiders in amber and copal. Conclusions, revisions, new taxa and family diagnoses of Editor: Brent Emerson

1266 Journal of Biogeography 41, 1255–1266 ª 2014 John Wiley & Sons Ltd 104 RESULTATS

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

124 RESULTATS

(& & '* ( '' ( ' ( ' &# #$# % ( & ', - ', , 0 7 R #8# &'( '( ( 7 A , - % (B A N A # && *( (6 * #

#P ! * m m " m & m m m m m m #

125 Article 3

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

126 RESULTATS

( ( 1; 89?; & #8#

$Q ! m m m " m % # &$% '()

+ ( &# ; 7;; ( :V# B * ( ' 8" # ( & 6 (' #

127 Article 3

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

128 RESULTATS

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

129 Article 3

%P ! * m m " m & m m m m #

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130 RESULTATS

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

131 Article 3

* ( # % &8# ( * ( ( & # ( ( ( ( & #6; &1 "#74 * &' ( & #

&Q ! m m m " m % # &$% '()

132 RESULTATS

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

133 Article 3

'P ! * m m " m #

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134 RESULTATS

' (0 ( ( ' ( & ( * (( (#

(Q ! m m m " m % # &$% '()

''' & (( ( #$# #7' #$# ( ' ( & & ((' ( *( ( & ' ( #! ' & (* ( * ( ' ( ' #:# ''' #$# 1 ' & ( - & ( (

135 Article 3

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

136 RESULTATS

#6; &1 "#,8 W"+""#86" #!" +""#:/" W"+"#"6"!#: +"?# !/ W"+8!#:6" #9 + 8:#?/ W"+" #!6""# : +"8#?/, W +"#9!6 #8" +"#! $ W +"#?"6"#?9 +"#? #

)P ! * m m " m #

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137 Article 3

9# H 8#88H 9# !H 9# H 8#" ,"8:#:"+ 9#?H8#8?H9#!H#8"H8# : , 8#9:+ 9# 8H8#8!H#"H9# 8H8#8 , 8!#+ 9#" H8# H#:9H9#:H8# #. #: ' * ( ( & #' * ' &* (4 # 7 ( & & * ( # ; % # * ( * ( # ; ( ( ( ( &(& ( ' ( #64 * &' ( & # 1 '# O * ' * 6 1 '#% '' * ( * ( ' ' ( #

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138 RESULTATS

0/?9 0/???1/?:90/! ?/!8 /!!"/: :1 ( ( 1; 89?; & #8#

* Q ! m m m " + m # &$% '()

+ ( &# ; ( ? V#,' * ( # &* (&( ( &4 # 4 564 & ( # 8 # * (

139 Article 3

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

140 RESULTATS

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

141 Article 3

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+ ( &# ; ( :V#,' * ( / , 8, 4 # &* (*( &( ( &4 # 4564 & ( #88#* ( ( 4 #& 4#( ( * 4 #;

142 RESULTATS

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

143 Article 3

P ! * m m " m #

. B &6,L8?!:# ; :V ( #0 * ( #( , , #; #&*6 * (( ( &4 #B ( (#456( ( '( ( # ( & #?8 ( * # 9#B( "#9" 8#89* #& -+, #"?B7 #8B, #"8#, '7 & #8?#; " ( * 8## 9#" #* *# , , ' 6# , 8" # :+ ' #9?H 8# "H #:?H #?"H 8#98 , " "9#"?+?#89H8# 9H?#?"H?#H8#9 , " #9!+ # 8H8# :H9#9H# 8H8# ?,6+6H6H6H6 . #8" ' * ( ( & # ' & * ' &* (4

144 RESULTATS

#7 ( & & * ( # ; % # * * ( #; * ( ( ( ( ( & ( ( ' ( #6; & '' ( ' ( #; &1 "#,8 W+8#996 "#9! +"8#""/" W9+8#86"?# +""#9/ W+8 #?!6"8#9 +8?#?9/ W9+89#?86"#!" +8:#:/, W9+"#!:6 #?: + #8/$ W9+"# 6 # ! + #8#0 '6&* 4&*6 ( ' ( #1((' (456 4 & % # ,0 3 # O * ' ( ( ',0*( (& ' ' 3 #

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145 Article 3

# mm m m+ mm m , m,!- m mm mm.'/ m

/ ? 1( " " / #$#/88 #$#/' #$#/ #$#/ #$# ' #$#3 & *4' ( "F #8# ' (( & *4'( *4% ' #$#*( ( * *4# ) "(*( (' '' * ( ( " ( & '( # ' (F * ( (* ' ' "

146 RESULTATS

$0 12 m m m mm 3 m mmm m m mm m m4 5 m m4 m m 3 % m m m m mm m06m

147 Article 3

' ( ( (& ( ##7A63 Z - " 8 #1 ' & ( * ( & & ( & ' / ('* 4& (% & "#

1( ') & 8" /" #$#": #$# % #$# # 18/ #$# * #$#8? #$# % #1(% ' (< = ( . * ( % ' ( ' * ' & #89# ' (8+8 ( ( % ' * ( ( ( ( # 1( ((' ( * * 9#9 :#?MO"B *( ( ' * M# ' & * * ( * '' + ()) (C, 8M ((#1()) 8 M ' & ( F ( ' &#& (C, '* ' 7&>B&" 9# *' ( * *' ( (; & #8#

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148 RESULTATS

(*(' '' ( (' ( #1( * ( '' ' * & ( *( & &' & ( ( 0 (8:9!/8:?/0 (> 48:! #& (* '' * * ( * ( *& ' ( 8" M B > " 8 ( ' < = # 89 ( ; & #8 ' & " ( & # 8# 1( ( (& ( ' ( & ( ( ' @ -" ?#

% mm m m m 7 m mm m 7 m * m mm 8 9 mm m m m - m m : m m m '' .;/7# m /

$ ' ' & # 168 ' 1 ' ( &B > " 8 # C & . ' &* ' ( / (' * & # (( ( (( 4 56(4 * ( ((&' ( ( )F (* ('' (F ' ( ' G ) G) ' ' * *" 8 /4 #" 8 * '&

149 Article 3

( * ' ( 1 '*( ( * ' # ) ' ' ( ( <4 % = B" 88 * ( F # ; ( ' ( & (& (' &' ' ( * ( & ' 1 ' '' & * # 1( ( (* ( (' # 168' (& (& & * ( 4 B > " 8# ( * & ( &( ( * ( ( '456(4 ( ' (( ( #1( ( & ## ( ( B > " 8/ ( (7 '1 %( 44 (( >B " :# ( 0 (8:9!8:?/ 0 (> 48:! '' ( (& (& & * ( ' #" !#0 ( (& & B > " 8 (( ' ## ( ( % * ( ( 0 (8:! ' #" !#

@ 7 & & ( ( &' ( (< 6=5 5 8:? #1( * ( & * ' **' ' # (& &(* ( ( (& ( *& 7 & >; '' " /7 >B " !/7 > " 8 / #" 8#1 4 * (< 6=( % & (( %' % ( ( 3* 3 " " ( 7 8::8 ( N Z - ( 8:"# ( & (( (&

150 RESULTATS

%' '% > 8::? ; 8!! '*' '% $ (8::" > 8::?#1( %* ( ( & #$# 1 '#1( ( & (* (* ( ' &( ( ( ( ( #8 # & ( & * % ' & #$# ' , - ( * ( (* ,H, , (H ( # $# ( * ( (( ( ,H, '* & # '* & ( & < 6='' ( * &( . ((& ( ' '1 ' ,0B > " 8 ( ',0 1 '#) ( ( %' ' ( *( (( ( ( (& ( #3* ( * ( ( ' ' # & ( ' ( / ( & ' ( *' * (* 7;; # & #$# ( & % & / ( (% & ( ( ( '' ' #

$ (( % # ( ( ' ' ( #) ( (* ' (' ' # " : & & % 4 * &' ' &#1( 4 * ' & ' & C,*( ( ' ' (F&F (* ( (* ( ( 8M

151 Article 3

' F (>3 " ?#$(* ( ( (( &' C, ##8M* ' &* ( ( & - (('9#9M' ' & #1( * ( B #" 88/5 #" 8" ( *( ( . ' () < &= ( ( < ' = > 4( 4" 8 #) ()6 ' ' & & * ( ' ' '% *( '& # & ) ( & '& 5 41(>BF " 9 ( # F & ( ') '& * &( ( 4 *' ( #

%)(,&5&+&! $ ( 4 7# & # # 0 -Z- # N *4 5# C *( ' *4 )#7AB#CA B#NX' ' ( &#$ ( 4C#$ ;# 65 ' # ' ( ( * & ( 0," !6 !9 ' ( 7 U & ; ##B#* & B '7 &1 A;6" :6 89!?8#

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152 RESULTATS

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153 Article 3

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154 RESULTATS

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

155 Article 3

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156 RESULTATS

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168 RESULTATS

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169 Article 4

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170 RESULTATS

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171 Article 4

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172 RESULTATS

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177 Article 5

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178 RESULTATS

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179 Article 5

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180 RESULTATS

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181 Article 5

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Diversitat del gènere Loxosceles al Nord d’Àfrica

Article 6 Ribera C, Planas E. 2009. A new species of Loxosceles (Araneae, Sicariidae) from Tunisia.

Article 7 Planas E, Ribera C. On the shoulders of Atlas: high genetic diversity of Loxosceles spiders (Araneae: Sicariidae) in the Souss-Massa and adjacent regions (NW Africa).

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A peer-reviewed open-access journal ZooKeys 16: 217-225 (2009) A new species of Loxosceles (Araneae, Sicariidae) 217 doi: 10.3897/zookeys.16.232 RESEARCH ARTICLE www.pensoftonline.net/zookeys Launched to accelerate biodiversity research

A new species of Loxosceles (Araneae, Sicariidae) from Tunisia

Carles Ribera†, Enric Planas‡

Institut de Recerca de la Biodiversitat de la Universitat de Barcelona (IRBio) and Departament de Biologia Animal, Facultat de Biologia, Universitat de Barcelona, Av. Diagonal, 645, 08028 – Barcelona, Spain

† urn:lsid:zoobank.org:author:509826D6-1E4D-4201-B08C-07D0F0044CC7 ‡ urn:lsid:zoobank.org:author:2387D04B-02FD-4884-8CF4-578A3A1DE067

Corresponding author: Carles Ribera ([email protected])

Academic editor: Jason Dunlop | Received 13 March 2009 | Accepted 11 May 2009 | Published 29 July 2009

urn:lsid:zoobank.org:pub:2DCADC67-CC80-4F81-95DC-0C1AF57A6371

Citation: Ribera C, Planas E (2009) A new species of Loxosceles (Araneae, Sicariidae) from Tunisia. In: Stoev P, Dunlop J, Lazarov S (Eds) A life caught in a spider's web. Papers in arachnology in honour of Christo Deltshev. ZooKeys 16: 217-225. doi: 10.3897/zookeys.16.232

Abstract A new species of the spider genus Loxosceles, L. mrazig sp. n., found in Tunisia is described and illustrated. Th e male bulb shows a high degree of morphological similarity to that of L. gaucho from Brazil, but the proportions of the palpal segments and the general colouration of the body reveal signifi cant diff erences between the two species. A distance analysis of the sequences of the mitochondrial gene cox1 reveals that the specimen from Tunisia shows high genetic distance from L. gaucho (more than 20%). Th e American species L. gaucho and L. laeta form a sister group to the Mediterranean representatives (L. rufescens and the Tunisian specimen).

Keywords Taxonomy, Araneae, Loxosceles, new species, Tunisia

Introduction Th e genus Loxosceles Heineken et Lowe, 1832 is currently known to comprise 97 species (Platnick 2009), 82 of which occur in America, 12 in Africa and two in China. Following Brignoli’s (1969, 1976) contributions with respect to the Mediterranean basin, only a single species is currently accepted as valid, L. rufescens (Dufour, 1820),

Copyright Carles Ribera, Enric Planas. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

209 Article 6

218 Carles Ribera & Enric Planas / ZooKeys 16: 217-225 (2009)

whose type locality is near Sagunt, Valencia (Spain). Th e other two (sub-) species are considered nomina dubia: L. decemnotata Franganillo, 1925 from Spain and L. rufescens lucifuga Simon, 1910 from Algeria. In the same paper Brignoli (1976) reported the South American species L. gaucho Gertsch, 1967 from Tunisia. In 2007 colleagues from the Ecology Department at the University of Barcelona collected in Douz (Tunisia) a male of Loxosceles in a dune located several kilometres from the city. Th e morphology of the copulatory bulb is remarkably similar to that of L. gaucho from Brazil, although the diff erences of the proportions of the male palpal segments plus the general colouration of the body suggested that it could be a diff erent species. In order to test this curious distribution, we used the cytochrome oxidase I gene (cox1) to compare this new record with L. gaucho (Sao Paulo, Brazil).

Material and methods Taxonomy. Specimens were examined under a Zeiss Stereo Discovery V12 stereomi- croscope equipped with an Infi nity X DeltaPix digital camera. Digital microscopic im- ages were edited using DeltaPix DpxWiew Pro AZ V. 13.6 software, using an enhanced focus function. Ink drawn digital illustrations were generated with the assistance of Photoshop CS3 software. Measurements were taken using the enhanced focus function incorporated into the DeltaPix DpxWiew Pro AZ software. All morphological measurements are given in millimetres. Prosoma and opisthosoma measurements were taken in dorsal view. Total body length represents the sum of the lengths of the prosoma and opisthosoma, omitting the pedicel. Eye largest diameters were taken from the spans of the lens. Th e largest leg article lengths were measured in lateral view without detaching the legs from the specimen, by placing the article being measured in a perpendicular position. Holo- type, and all other specimens are deposited in the Arachnid Collection of the CRBA (Centre de Recursos de Biodiversitat Animal) at the University of Barcelona; catalogue numbers are given in brackets. Abbreviations used in the text. CRBA – Centre de Recursos de Biodiversitat Animal, Universitat de Barcelona, Spain. Eyes: ME = median eyes; LE = lateral eyes. Molecular data. Taxonomic sampling. Taxa analyzed in the present study are listed in Table 2. L. mrazig sp. n. from Douz (Tunisia), L. gaucho from Sao Paulo (Bra- zil) and nine specimens of L. rufescens from diff erent localities in the Iberian Peninsula and Tunisia were analyzed. In addition we included a representative of Loxosceles laeta (Nicolet, 1849) (Montevideo, Uruguay) in order to test the phylogenetic affi nities of L. gaucho with other South American species, since L. laeta belongs to a diff erent species group (Gertsch 1967; Binford et al. 2008). A sequence from Dysdera crocata C. L. Koch, 1838 from GenBank was also included to root the tree. Sample Storage and DNA Extraction. Specimens were preserved in 95% or absolute ethanol and stored at 4°C. Total genomic DNA was extracted from legs of a single specimen using the QIamp® DNA Mini Kit (QIAGEN) following the manufacturer’s

210 RESULTATS

A new species of Loxosceles (Araneae, Sicariidae) 219 protocols. Th e approximate concentration and purity of the DNA obtained were veri- fi ed using 1.5% agarose/TBE gel electrophoresis. PCR Amplifi cation and Sequencing. A total of 899 bp of the cytochrome oxidase I gene (cox1) was amplifi ed from each individual using PCR with the following primer pairs: C1-J-1718 (Simon et al. 1994) with C1-N-2776 (Hedin and Maddison 2001). Th e PCR reaction mixture contained a fi nal concentration of 0.2 μM of each primer, 0.2 mM of each dNTPs, 0.5 U Taq polymerase (Promega), with the supplied buff er, and 1.5-2.5 mM Mg Cl2 in a fi nal volume of 25 μL. A Perking-ElmerCetus Moldel 480 thermocycler was used to perform 35 iterations of the following cycle: 30s at 94°C, 45s at 44°C, and 1 min at 72°C, beginning with an additional step of 3 min at 94°C, and ending with another step of 5 min at 72°C. Th e PCR results were visualized by means of a 1.5% agarose/TBE gel. Amplifi ed products were purifi ed using MultiScreen 96 – well fi lter plates from Millipore. Th e purifi ed products were directly cycle-sequenced from both strands using ABI BigDye (Applied Biosystems) chemistry and run out on ABI Prism 377 (Applied Biosystems) automated sequencers. Sequencing reactions were performed in our lab with the forward and reverse PCR primers and one additional pair of internal cox1 primers, CI-J-2183 and C1-N-2191 (Simon et al. 1994). Th e resulting products were run and analyzed at the Serveis Científi co-Tècnics of the Universitat de Barcelona. Alignment. Raw sequences were compared against chromatograms and comple- mentary contigs built and edited using the Geneious Pro 3.6.2 software (http://www. genious.com). Sequences were manipulated and preliminary manual alignments con- structed using BioEdit V.7.0.5.3 (Hall 1999). Alignment of cox1 was trivial, given that no evidence of insertions/deletions was observed. Genetic distances and distance analyses. Uncorrected genetic distances between and within taxa were estimated with MEGA v.3.0 (Kumar et al. 2004). Th e Neigh- bour-joining algorithm was applied to the estimated genetic distances to build a phe- nogram (Saitou and Nei 1987) conducted with the same program. Clade support was assessed via Bootstrap (Felsenstein 1985) as implemented in MEGA, based on 1000 bootstrap replicates. Results. Molecular data. Th e distance tree is shown in Fig. 1. Th e specimens iden- tifi ed as L. rufescens form a monophyletic clade with a high support value (100%). L. mrazig sp. n. is supported as more closely related to L. rufescens (75% bootstrap value) than to the two South American representatives included in this analysis: L. gaucho and L. laeta. Th e latter two species cluster together with moderate support (68%). Averages between group genetic distances are presented in Table 1. Th e cluster formed by the nine specimens of L. rufescens from Spain and Tunisia shows scarce

Table 1. Average between group genetic distances of gene cox1 from the four species analyzed. L. rufescens L. mrazig L. gaucho L. mrazig 0.1991 L. gaucho 0.1927 0.2063 L. laeta 0.1973 0.2086 0.1635

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220 Carles Ribera & Enric Planas / ZooKeys 16: 217-225 (2009)

within-group average genetic distances (0.26%) and suggests that this species shows a high genetic coherence. Th e deep genetic divergence between L. mrazig and L. gaucho (20.63%) together with the observation that both species belong to diff erent clusters provide clear evidence that L. mrazig is an independent evolutionary lineage and should, therefore, be considered a diff erent species.

Table 2. Species included in the phylogenetic analysis and GenBank accession numbers for cox1. Species Locality GenBank Accession number Loxosceles laeta Montevideo, Uruguay FJ986177 Loxosceles gaucho Sao Paulo, Brazil FJ986178 Loxosceles mrazig sp. n. Douz, Tunisia FJ986179 Loxosceles rufescens Torrejon de Ardoz, Madrid, Spain FJ986183 Loxosceles rufescens Ciudad Real, Spain FJ986185 Loxosceles rufescens Denia, Alacant, Spain FJ986187 Loxosceles rufescens Chumilla, Murcia, Spain FJ986181 Loxosceles rufescens Barcelona, Spain FJ986182 Loxosceles rufescens Siles, Jaen, Spain FJ986188 Loxosceles rufescens Sierra Gorda, Cartagena, Murcia, Spain FJ986180 Loxosceles rufescens Alacant, Spain FJ986184 Loxosceles rufescens Testour, Tunisia FJ986186 Dysdera crocata Hoz de Pergrina, Guadalajara, Spain EF458137

Torrejon Testour CReal Denia Chumilla L. rufescens BCN Siles 100 Cartagena 75 Alicante L. mrazig sp. n. 68 L. laeta L. gaucho Dysdera crocata

0.02 Figure 1. Neighbour-joining distance tree. Diff erent representatives of L. rufescens from the western Mediterranean basin (Spain and Tunisia), L. mrazig sp. n., L. gaucho and L. laeta are included. Numbers on nodes represent bootstrap support values.

212 RESULTATS

A new species of Loxosceles (Araneae, Sicariidae) 221

Taxonomy Family Sicariidae Genus Loxosceles Heineken et Lowe, 1832

Loxosceles mrazig sp. n. urn:lsid:zoobank.org:act:A8F75878-85CA-4567-874B-EBC8CC24CD02 Figs 2-7

Material examined. 1 male (Holotype) from Douz, Tunisia, 33° 24’ 26.77’’ N, 09°02’41.92’’E, 27 January 2007, Cesc Múrria leg. (CRBA-LX1054). Material for comparison. 2 males, 2 females of L . gaucho (CRBA-LX1024) from Sao Paulo, Brazil, November 2007, A. Brescovit leg.; 2 males, 2 females of L. laeta (CRBA-LX1028) from Montevideo, Uruguay, L. Acosta leg.; 1 male, of L. rufescens (CRBA-LX1012) from Gavà, Barcelona, López-Pancorbo leg. Etymology. Th e species’ name honours the people called Mrazig, formerly nomad- ic, living in and around the city of Douz (Tunisia). Th e Mrazig are the descendants of the Banu Saleim tribe that fl ed the Arabian Peninsula in the seventh century and came to Tunisia in the thirteenth century. It is known that they practiced transhumance in the Great Sahara. Noun in apposition. Diagnosis. Diff ers from L. gaucho, L. rufescens and its similar relatives in the pro- portion of male palp segments, mainly the tibia. In L. mrazig the tibia is markedly oval, slightly longer than wide (0.63 - 0.54) (Figs 2, 3, 5); in L. gaucho it is ¾ as wide as long (Gertsch 1967, plates 3-4), whereas, in L. rufescens, it is slightly oval, although dorsally almost straight (Gertsch 1967, plate 10). Also diff ers from L. rufescens by the size of the tegulum and the size and shape of the embolus (Figs 2-5). Body pigmentation yellowish-brown in L. gaucho and pale yellow in L. mrazig. In general, the morphological diff erences compared to L. rufescens are more conspicuous. Th e size of the tegulum and, especially, the shape and length of the embolus are clearly diff erent. Description. Colouration: Carapace pale yellowish with a fi ne, pale brown lateral stripe. Median groove and adjacent integuments darkened. Pars cephalica slightly darkened, brown coloured, and clearly demarcated by a lateral reddish brown line. Less conspicuous, but still important, diagnostic traits are the four thin longitudinal lines (lightly impressed when seen under higher magnifi cation) located in the centre of the pars cephalica (Fig. 6). Eye tubercles black. Sternum pale yellowish, paler than carapace. Labium and gnathocoxae with slightly more pigmentation. Legs light yellow or some- what shaded, with the apical segments slightly darkened. Opisthosoma yellowish-white. Prosoma. Carapace (Fig. 6) slightly longer (2.39) than wide (2.15). Median groove deep, occupying the posterior third of carapace. Clypeal width slightly more than 2.5 diameters of ME. Eyes close together (Fig. 7); LE separated from ME by the diameter of ME. LE larger than ME (0.18 - 0.1 respectively). Sternum about ⅔ as wide as long, extended between the IV pair of coxae. Labium as long as wide at its base, apically nar- rowed and rounded. Gnathocoxae distally convergent, enclosing the labium.

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222 Carles Ribera & Enric Planas / ZooKeys 16: 217-225 (2009) 0.5 mm

230.5 mm

4 5

0.5 mm 0.5 mm

Figures 2-5. Male palp of Loxosceles mrazig sp. n. 2 prolateral view 3 retrolateral view 4 apical view 5 dorsal view.

Opisthosoma elongate oval in dorsal view. Male palp (Figs 2-5). Femur cylindrical, more than fi ve times longer than wide. Tibia short, oval, slightly longer than wide. Tarsus fl attened below, slightly shorter than tibia, rounded apically. Tegulum large, 4/5 as wide as tarsal length. Embolus enlarged at base, forming a sinuous curve, about 1.5 times longer than tegulum. Measurements. Male (holotype): Prosoma 2.15 wide, 2.39 long: opisthosoma 3.22 long. Total body length 5.61. Legs: I: coxa 0.81, trochanter 0.23, femur 5.42, patella 0.84, tibia 5.70, metatarsus 5.76, tarsus 1.38, total length 20.14; II: coxa 0.58, rest of segments missing. III: coxa 0.81, trochanter 0.23, femur 4.94, patella 0.82, tibia 4.69, metatarsus 5.37, tarsus 1.12, total length 17.98; IV: coxa 0.81, trochanter 0.23, femur 5.26, patella 0.84, tibia 5.40, metatarsus 6.42, tarsus 1.33, total length 20.29; Palp: femur 1.19, patella 0.36, tibia 0.63, tarsus 0.56, total length 2.74. Female unknown. Distribution. So far, L. mrazig is known only from the type locality. Th e unique specimen was collected in a dune of sand near the city of Douz.

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A new species of Loxosceles (Araneae, Sicariidae) 223

1 mm

Figures 6-7. Prosoma of Loxosceles mrazig sp. n. 6 dorsal view, arrow indicates the four longitudinal lines located in the centre of the pars cephalica 7 frontal view.

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224 Carles Ribera & Enric Planas / ZooKeys 16: 217-225 (2009)

Discussion Th e possibility that this species should be assigned to L. gaucho can be ruled out due to the high genetic distance observed between both species (more than 20%) and especially because they do not form a sister group relationship, but belonging to diff erent clades. Th e morphological similarity can be explained as a convergence phenomenon due to the simple morphological structures of the copulatory organs found in haplo- gyne spiders. Determining the closest relatives is diffi cult for this species due to the lack of current knowledge on African Loxosceles species. Taking into account the shape of the male bulb, this species could be related to L. foutadjalloni Millot, 1941 from Guinea, in which the proportional palpal segments diff ers notably (mainly the tibia) and by the shape and size of the embolus. L. mrazig sp. n. – which could possibly be a member of a diff erent group, or form a subgroup with the above mentioned L. foutadjalloni. L. mrazig sp. n. – is the second Loxosceles species known from the Mediterranean basin.

Acknowledgements We would like to thank Luís Acosta from Uruguay and Antonio Brescovit from Brazil for their inestimable help and collaboration in providing us with the specimens of L. gaucho and L. laeta for this work. We also thank Cesc Múrria for collecting the new species and Miquel A. Arnedo for providing the sequence of D. crocata. Th is research was supported by the Spanish Ministry of Education and Science grants CGL2008- 03385/BOS and CGL2006-13374/BOS.

References

Binford GJ, Callahan MS, Bodner MR, Rynerson MR, Berea Núñez P, Ellison CE, Duncan RP (2008) Phylogenetic relationships of Loxosceles and Sicarius spiders are consistent with Western Gondwanan vicariance. Molecular Phylogenetics and Evolution 49: 538-553. Brignoli PM (1969) Note sugli Scytodidae d’Italia e Malta (Araneae). Fragmenta entomologica 6: 121-166. Brignoli PM (1976) Beiträge zur Kenntnis der Scytodidae (Araneae). Revue suisse de Zoologie 83: 125-191. Felsenstein J (1985) Confi dence limits on phylogenies: an approach using the bootstrap. Evolu- tion 39: 783-791. Gertsch WJ (1967) Th e spider genus Loxosceles in South America (Araneae, Scytodidae). Bul- letin of the American Museum of Natural History 136: 119-173. Hedin MC, Maddison WP (2001) A combined molecular approach to phylogeny of the jumping spider subfamily Dendryphantinae (Araneae: Salticidae). Molecular Phylogenetics and Evolution 18: 386-403.

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A new species of Loxosceles (Araneae, Sicariidae) 225

Hall TA (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95-98. Kumar S, Tamura K, Nei M (2004) MEGA3: Integrated software for molecular evolutionary genetics analysis and sequence alignment. Briefi ngs in Bioinformatics 5: 150-163. Millot J (1941) Les araignées de l’Afrique Occidentale Française - sicariides et pholcidés. Mé- moires de l’Académie des Sciences de l’Institut de France 64: 1-53. Platnick NI (2009) Th e world spider catalog, v. 9.5. American Museum of Natural History, online at http://research.amnh.org/entomology/spiders/catalog/ index.html Saitou N, Nei M (1987) Th e neighbor-joining method: a new method for reconstructing phylogenetic trees. Molecular Biology and Evolution 4: 406-425. Simon C, Frati F, Beckenbach A, Crespi B, Liu H, Flook P (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America 87: 651-701.

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228 RESULTATS

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229 Article 7

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230 RESULTATS

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231 Article 7

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232 RESULTATS

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233 Article 7

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234 RESULTATS

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235 Article 7

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236 RESULTATS

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237 Article 7

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238 RESULTATS

$# ## " " " " " $!! $ " " " " " $$ !$# " " " " " $# # " " " " " !! $$ " " " " " ! !!!$ %&##!! %&##!# %&# " %&##! ) $! !$ $ !# " " " " " ) $ ! " " " " " 9 $#! ! &*# +## %&#$ " %&##!# ,-./0 !$ # #! " " " " ,-./0 ! $$! " " " " " ,-./0 !$ ##! " " " " .&1/2/3/12 # %& %&#!! $ %&#!!## " %&#! 42/ . 22 #! $$ %& %&#!!# " %&#!! . 22 ! $$# %&# %&#!!$# %&#!!#! " %&#!! . -/2 !## #$ %&#!!! %&#!! %&#!! %&#! . //2/ # $# # %&$ %&#!! %&#!!# " %&#!! . //2/ $$ $ %& %&#!! %&#!!# " %&#!!#

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CAPÍTOL 4

Diversitat del verí i loxoscelisme a la Conca Mediterrània i a les Illes Canàries

Article 8 Planas E, Zobel-Thropp PA, Ribera C, Binford G. Not as docile as it looks: Loxosceles venom variation and loxoscelism in the Mediterranean Basin and the Canary Islands.

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245 Article 8

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246 RESULTATS

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247 Article 8

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248 RESULTATS

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249 Article 8

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250 RESULTATS

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251 Article 8

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252 RESULTATS

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254 RESULTATS

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255 Article 8

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256 RESULTATS

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258 RESULTATS

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259 Article 8

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260 RESULTATS

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261 Article 8

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262

DISCUSSIÓ GENERAL

)

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3655 * ) 655

267 * # , ( 3 347 + ) ' # 8 + ' 8 ) * /( 60 * 9 : 1 , # . ) & ) + # 1 ) & /; # 653<0# (& ( ,& ) - # ( - ; /3=>?0# (& & * #

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268 DISCUSSIÓ GENERAL

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270 DISCUSSIÓ GENERAL

( 3 / B 0# +

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271 + ) & )

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272 DISCUSSIÓ GENERAL

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274 DISCUSSIÓ GENERAL

+ .B + '

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276 DISCUSSIÓ GENERAL

) & ) # , ) ) ) / 8 ; 3=>?0# , + & '

+ /V 65550# ( 7 - + *

B + G3 G G , G 2 # K & D + # 8

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277 ;6 /(? ;<0 + ) / L > +0 + ) / L C ( 30 ( 3 ) +

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278

CONCLUSIONS

!" # $ %&' (!)" * #

) + , $ '( %- . !/#" 0 $ ' ( 1 0 (

2 + * #

3 + ' - ' ' & * ( 4 1

5 + , ( - 6*7 ( ' / 8 ( , 9 7 -.( : 6*

; + ' 4 ' - '*

< + . ' =' ( ' ' > (+ ( , ? ' 6 *?

283 ' 7 ? ' +6 / @ 7 '

A / ' 8 ' & * B

C '1 > D + ( = ( ' . '1 B

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+ ' 4 > ( .+ ( $ '

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2 + ' B ' D# 1 - . .

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284 CONCLUSIONS

' & * 78 ' ' - % ' ( ' * B

; + ' * 8 ' ' / % ' ' & * ' ( ( '

285

CONCLUSIONS

F !" # $ % &

#& ' ' ( " $ " &

)& * " % &

+& ' , , " - " &

.& / " , 0- / &/ 1 " " 2 &3 45 " " 0- &

6& / " - &

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287 &

:& " " - ! " " &

;& < $ " 3 81 , 5 &

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.& / << '0( " " - ! & " ,

288 CONCLUSIONS

, - " &

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289

REFERÈNCIES

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294 REFERÈNCIES

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296 REFERÈNCIES

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298 REFERÈNCIES

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