A revised classification of Santalales

Abstract

We present here a revised classification of Santalales, an angiosperm order that contains 18 families, 160 genera, and over 2200 species. Both nonparasitic and parasitic flowering plants occur in the traditionally circumscribed family Olacaceae whereas all other families are composed entirely of parasites. The five evolutionary radiations of aerial parasitism produced mistletoes that constitute most of the generic and specific diversity seen in the order. This classification, although based primarily upon results from molecular phylogenetic investigations, brings together all currently available information that contributes to our understanding of relationships among these plants. Monophyletic groups (clades) obtained from molecular analyses were named using a Linnaean ranked system. Four new families are named that formerly resided in Santalaceae s.l.: Amphorogynaceae, Cervantesiaceae, Comandraceae, and Nanodeaceae. A new tribal and subtribal classification for Loranthaceae is presented where nine new subtribe names are proposed.

Full text

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INTRODUCTION
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TRADITIONAL TAXONOMIC TREATMENTS
OF SANTALALES
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A r e v i s e d c l a s s i f i c a t i o n o f S a n t a l a l e s
Daniel L. Nickrent,1 Valéry Malécot,2 Romina Vidal-Russell3 & Joshua P. Der4
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Abstract
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Keywords
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TA XO N O MY

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TA XO N$!%$&'($)$*+,-.$'/0/1$!"#2!!#3-45,678$9$:.;$)$Classif ication of Santalales
9$3-45,678$&'//#?($>=@F6B$8=:8$ J":2C#%C)80$F:>$8=6$E-,>8$
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FEATURES CHARACTERIZING SANTALALES
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8F@$=:A6$?667$.@>8$.6:A-7D$8=,66$>8:C-7@B6>;$<7$8=6$!OC6,O
@L>$NH$ :+&%C#%:$]@S?;$8=,66$ >8:C67>$:,6$E6,8-.6$ :7B$E-A6$:,6$
>8:C-7@B6>;$T@,6@A6,J$8F@$@E$ 8=6$ 8=,66$ E6,8-.6$ >8:C67>$:,6$
:786>6+:.@L>$ :7B$@76$@44L,>$@7$8=6$@++@>-86$E .:75$@E$:$+68:.$
:.@7D$F-8=$:$>8:C-7@B6;$K=->$+68:.$:.>@$=:>$8F@$A:>4L.:,$8,:46>$
-7>86:B$@E$@76;$K=6>6$@?>6,A:8-@7>J$+.L>$8=6$E:48$8=:8$>@C6$N*&_$
:,6$=6S:C6,@L>J$4@C+6.>$@76$8@$4@7>-B6,$ 8=:8$ 8=6$E.@,:.$+.:7$
@E$+678:C6,@L>$>+64-6>$->$+>6LB@B-+.@>86C@7@L>J$-;6;J$B6,-A6B$
E,@C$:$8,-C6,@L>$+,@8G+6;$K=->$+.:7$->$.-56.G$8=6$@76$E@..@F6B$
?G$`:,G@+=G..:.6>$&]@7>6$M6$`,:676$9$:.;J$0%%#(J$F=-4=$E,@C$
,64678$C@.64L.:,$ +=G.@D6768-4$ 6A-B6746$&T@@,6$9$:.;J$'//#b$
Q@.8->$9$:.;J$'//"($->$8=6$76S8$4.:B6$8@$6A@.A6$:C@7D$8=6$:>86,-B>$
:E86,$Q:78:.:.6>;
HL?.->=6B$4=,@C@>@C6$7LC?6,>$=:A6$?667$4@C+-.6B$E@,$
:..$Q:78:.:.6>$D676,:$6S:C-76B$&Y.648,@7-4$QL++.6C678$8@$8=->$
:,8-4.6(;
CLASSIFICATION PHILOSOPHY
<7$8=->$ +:+6,$F6$+,6>678$ :$ ,6A->6B$4.:>>-E-4:8-@7$>G>86C$
E@,$Q:78:.:.6>$8=:8$-74@,+@,:86>$C@.64L.:,$:7B$C@,+=@.@D-4:.$
B:8:;$K=->$4.:>>-E-4:8-@7$->$766B6B$?64:L>6$,6.:8-@7>=-+>$:C@7D$
:..$C6C?6,>$ @E$8=6$ @,B6,$=:A6$7@8$?667$ :BB,6>>6B$ E@,$ C:7G$
B64:B6>$:7B$7@76$=:A6$L>6B$6S+.-4-8$+=G.@D6768-4$+,-74-+.6>;$
K=6$C:P@,$E6:8L,6>$@E$@L,$4.:>>-E-4:8-@7$>G>86C$:,6$?:>6B$L+@7$
C@.64L.:,$+=G.@D6768-4$8,66>J$C:-7.G$?64:L>6$4.:B->8-4$:7:.G>6>$
@E$C@,+=@.@D-4:.$4=:,:486,>$:,6$.:45-7D$E@,$C@>8$D,@L+>$ &8=6$
6S46+8-@7$?6-7D$[.:4:46:6$?G$T:.c4@8$9$:.;J$'//^(;$<7$:BB-8-@7$
8@$,64@D7-R-7D$C@7@+=G.68-4$D,@L+>J$F6$=:A6$4=@>67$8@$:>>-D7$
U-77:6:7$,:75>$8@$A:,-@L>$4.:B6>$F-8=-7$8=6$@,B6,$&a,6LB67O
>86-7J$'//!(;$V64:L>6$@E$-8>$>-R6J$F6$=:A6$:.>@$+,@+@>6B$-7E,:E:O
C-.-:.$,:75>$E@,$8=6$.:,D6$E:C-.G$U@,:78=:46:6;$a@,$E:C-.-6>J$F6$
=:A6$E@..@F6B$]64@CC67B:8-@7$0WV;0$@E$8=6$9A3S$&T436-..$
9$:.;J$'//!($8=:8$:BA->6>$:L8=@,>$8@$L>6$8=6$@.B6>8$7:C6;$K=6>6$
7:C6>$F6,6$@?8:-76B$+,-C:,-.G$E,@C$]6A6:.$&0%%X2($:7B$E,@C$
Z@@D.:7B$9$]6A6:.$&'//!(;
*.8=@LD=$F6$4@7>-B6,$8=6$:>>-D7C678$@E$,:75>$8@$?6$:$>@C6O
F=:8$>L?P648-A6$+,@46>>J$ F6$ :.>@$ B@$ 7@8$ E66.$ 8=:8$:$,:75OE,66$
>G>86C$&>L4=$:>$-,5*2A2C#($F-..$?6$EL..G$:446+86B$7@,$F-B6.G$
:B@+86B$-7$ :4:B6C-4$:7B$7@7:4:B6C-4$4-,4.6>;$_6$:457@F.O
6BD6$8=:8$ 8=6$U-77:6:7$ =-6,:,4=G$=:>$ :$ 7LC?6,$ @E$-7=6,678$
+,@?.6C>$>L4=$:>$&0($-7>LEE-4-678$,:75>$8@$674@C+:>>$:..$4.:B6>$
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:4,@>>$B-EE6,678$8:S@7@C-4$D,@L+>b$:7B$&"($C:7B:86B$7:C6$
4=:7D6>$8=:8$4:>4:B6$-78@$@8=6,$+:,8>$@E$8=6$4.:>>-E-4:8-@7$F=67$
8:S:$:,6$,6:>>-D76B$&H677->-J$'//0b$_-8=D@88J$'///(;$K=6$E-,>8$
+,@?.6C$->$7@8$:7$->>L6$-E$@76$:446+8>$8=:8$7@8$:..$4.:B6>$766B$8@$
?6$7:C6B$&*HIJ$0%%#(;$K=6$>64@7B$+,@?.6C$F@L.B$,6C:-7$6A67$
-E$,:75>$B-B$7@8$6S->8b$6;D;J$-7E@,C:.$7:C6>$L>6B$E@,$4.:B6>$4@L.B$
>8-..$?6$L>6B$-7:++,@+,-:86.GJ$E@,$6S:C+.6$F=67$4@C+:,-7D$8=6$
6S8678$@E$?-@B-A6,>-8G$:4,@>>$B->8:78.G$,6.:86B$8:S:;$*.8=@LD=$F6$
4:7$>6.B@C$?6$46,8:-7$8=:8$:$+:,8-4L.:,$+=G.@D6768-4$,6.:8-@7>=-+$
->$57@F7J$8=6$8=-,B$+,@?.6C$?64@C6>$.6>>$@E$:7$->>L6$:>$4.:B6$
C6C?6,>=-+$->$>8:?-.-R6B$F-8=$-74,6:>-7D$:C@L78>$@E$B:8:;$[L,$
4.:>>-E-4:8-@7$E@..@F>$8=6$7@C674.:8L,:.$+=-.@>@+=G$B6>4,-?6B$
:>$e8=6$C@7@+=G.68-4$U-77:6:7$+@>-8-@7f$&Q86A67>J$'//W(;
`.:>>-E-4:8-@7>$ ?64@C6$C@>8$L>6EL.$F=67$8=6G$ >-CL.8:O
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+6,>+648-A6$-E$8=6G$:,6$ C:S-C:..G$+,6B-48-A6$E@,$:..$ ?-@.@D-O
4:.$B->4-+.-76>$ :7B$ 8,L6$ E,@C$:$+,:48-4:.$ +6,>+648-A6$-E$ 8=6G$
>6,A6$ :>$ :$ C6:7>$ 8@$ @,D:7-R6$ :7B$ ,68,-6A6$ -7E@,C:8-@7$ &6;D;J$
=6,?:,-:J$E.@,:>J$I67V:75J$684;(;$VL8$:>$+@-786B$@L8$?G$QC-8=$9$
:.;$&'//W(J$8,:B-8-@7:.$:7B$C@.64L.:,$+=G.@D6768-4O?:>6B$4.:>O
>-E-4:8-@7>$:,6$@E867$-74@C+:8-?.6$-E$8=6$+,-74-+.6$@E$C@7@+=G.G$
->$:B@+86B;$K=6G$.->8$8=,66$B-,648-@7>$8=:8$4:7$?6$8:567$F=67$
:$8,:B-8-@7:..G$,64@D7-R6B$E:C-.G$,6>-B6>$F-8=-7$:7@8=6,$&-;6;J$
+:,:+=G.G(1$&0($,64@D7-8-@7$@E$+:,:+=G.68-4$ E:C-.-6>J$&'($>+.-8O
8-7D$8@$C:-78:-7$C@7@+=G.68-4$E:C-.-6>J$:7B$&"($.LC+-7D$@76$
E:C-.G$-78@$:7@8=6,;$_6$:D,66$8=:8$4=@-46$0$->$L7B6>-,:?.6$:7B$
:457@F.6BD6$8=:8$+-45-7D$?68F667$'$:7B$"$B6+67B>$+:,8.G$L+@7$
8:>86;$*.8=@LD=$8=6$.LC+-7D$:++,@:4=$+,@A-B6>$8=6$eNL-45$E-SJf$
B-A6,>-8G$F-8=-7$8=6$,6>L.8-7D$E:C-.G$F@L.B$.-56.G$,6>L.8$-7$8=6$
7:C-7D$@E$:BB-8-@7:.J$F6..OC:,56B$4.:B6>;
`,-86,-:$L>6B$8@$B64-B6$=@F$8@$4-,4LC>4,-?6$E:C-.-6>$E:..$
-78@$>6A6,:.$4:86D@,-6>J$?L8$:$?6D-77-7D$+@-78$E@,$B->4L>>-@7$
4:7$?6$8:567$E,@C$ V:45.L7B$9$V,6C6,$ &0%%#(;$*>-B6$ E,@C$
8=6$+,-C:,G$+,-74-+.6$@E$C@7@+=G.GJ$8=6G$.->8$ E@L,$>64@7B:, G$
+,-74-+.6>$8=:8$-7A@.A6$C:S-C-R-7D$&0($>8:?-.-8G$&j$C-7-C-R-7D$
7@C674.:8L,:.$4=:7D6>(J$&'($+=G.@D6768-4$-7E@,C:8-@7$&j$C-7-O
C-R-7D$,6BL7B:74G(J$&"($>L++@,8$E@,$C@7@+=G.GJ$:7B$&^($6:>6$@E$
-B678-E-4:8-@7$&j$,64@D7-R:?-.-8G(;$Q86A67>$&+6,>;$4@CC;($:BB>$
8F@$C@,6$+,-74-+.6>1$&!($+,6>6,A:8-@7$@E$D,@L+>$8=:8$:,6$F6..O
6>8:?.->=6B$-7$8=6$.-86,:8L,6$:7B$&W($E:C-.G$>-R6$@+8-C-R:8-@7;$
*886C+8>$8@$:B=6,6$8@$8=6>6$+,-74-+.6>$F6,6$C:B6$-7$8=->$F@,5$
:7B$>+64-E-4>$,6.:8-7D$8@$+:,8-4L.:,$B64->-@7>$F-..$?6$B->4L>>6B$
?6.@F$L7B6,$eH6,>+648-A6>$@7$8=6$36F$Q:78:.:.6>$`.:>>-E-4:O
8-@7f;
CLASSIFICATION OF SANTALALES
K=6$E@..@F-7D$,64.:>>-E-4:8-@7$@E$8=6$ C@7@+=G.68-4$@,B6,$
Q:78:.:.6>$4@78:-7>$0#$E:C-.-6>J$ 0W/$D676,:J$ :7B$@A6,$''//$
>+64-6>;$V:.:7@+=@,:46:6$:,6$ 7@8$ -74.LB6B$:8$ 8=->$8-C6;$ K=->$
4.:>>-E-4:8-@7$->$>LCC:,-R6B$-7$8=6$4@7>67>L>$+=G.@D67G$>=@F7$
-7$a-D;$0;$V64:L>6$@E$-8>$>-R6J$U@,:78=:46:6$:,6$EL,8=6,$4.:>>-E-6B$
-78@$8,-?6>$:7B$>L?8,-?6>$&a-D;$'$@7$+;$!^!(;
. Family Erythropalaceae Planch. ex Miq. (). —
$K=,66$
D676,:1$ O)5',)21&*80$ V.LC6$ F-8=$@76$>+64-6>J$ OH$:+&%C#%:$
V.LC6J$@E$<7B@C:.:G>-:b$M#":'#)"&$g:4N;$F-8=$4:;$""$>+64-6>$@E$
8,@+-4:.$*C6,-4:$:7B$8=,66$*E,-4:7$>+64-6>b$:7B$J&B8)#&$T::>$
F-8=$@76$>+64-6>J$JH$')"%#)<":$T::>J$@E$ILG:7:;
O)5',)21&*80$ ->$ :$ F@@BG$ 4.-C?6,$F-8=$ :S-..:,G$867B,-.>$
F=6,6:>$8=6$@8=6,$8F@$D676,:$:,6$>=,L?>$@,$8,66>;$V64:L>6$@E$
-8>$B-A6,D678$=:?-8J$8=6$:EE-7-8-6>$@E$O)5',)21&*80$=:A6$76A6,$
?667$4.6:,$:7B$=:A6$.6B$C:7G$F@,56,>$8@$->@.:86$O)5',)21&*80$
-7$-8>$ @F7$ E:C-.GJ$Y,G8=,@+:.:46:6;$ H.6>-@C@,+=-4$E6:8L,6>$
>=:,6B$?G$:..$8=,66$D676,:$:,6$:.86,7:86J$>-C+.6J$+68-@.:86J$6SO
>8-+L.:86$.6:A6>J$?->6SL:.$E.@F6,>J$.:45$@E$:$D.:7BL.:,$B->5J$:7B$

!^'
TA XO N$!%$&'($)$*+,-.$'/0/1$!"#2!!#3-45,678$9$:.;$)$Classif ication of Santalales
=G+@DG7@L>$@A:,-6>$4@C+@>6B$@E$8=,66$4:,+6.>;$K=6$+678:CO
6,@L>$E.@F6,>$C:G$?6$ B-+.@>86C@7@L>$ @,$->@>86C@7@L>$ F-8=$
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&:1*8%C""$Q.6LC6,($:7B$ 8=6$ E ,L-8>$ =:A6$ 8=-7$ 4,L>8:46@L>$67O
B@4:,+>;$ K=6$ :44,6>4678$4:.GS$ >@$4=:,:486,->8-4$ @E$M#":'#)"&$
->$7@8$ +,6>678$-7$8=6$@8=6,$8F@$D676,:;$ <7$8=6$C:8L,6$ E,L-8$ @E$
O)5',)21&*80J$8=6$?,-D=8$,6B$C6>@4:,+$>+.-8>$-78@$E-A6$,6E.6S6B$
A:.A6>$8=:8$,6>6C?.6$8=6$:44,6>4678$4:.GS$@E$M#":'#)"&$:7B$C:G$
>6,A6$:$>-C-.:,$EL748-@7$-7$:88,:48-7D$E,L-8OB->+6,>-7D$?-,B>;
K=6$ A:,-@L>$ :7:8@CGO$ @,$ C@,+=@.@DGO?:>6B$ >8LB-6>$ @E$
[.:4:46:6$=:A6$76A6,$4.6:,.G$-B678-EG$:$D,@L+$4@7>->8-7D$@E$D67O
6,:$-7$8=->$4.:B6$F=6,6:>$-8$,646-A6B$>8,@7D$>L++@,8$F-8=$V:G6>O
-:7$:7:.G>6>$ @E$ C@.64L.:,$B:8:$ &T:.c4@8$ 9$ 3-45,678J$'//#(;$
T::>$9$:.;$&0%%'($>8:86B$8=:8$F@@B$:7:8@CG$.-75>$J&B8)#&$8@$
M#":'#)"&$ F=6,6:>$+:.G7@.@DG$ &U@?,6:LO`:..67J$0%#/J$0%#'($
:7B$.6:E$:7:8@CG$ &V::>J$ 0%#'($.-75$-8$8@$8,-?6$`@L.6:6$ &=6,6$
E:C-.G$`@L.:46:6(;
. Family Strombosiaceae Tiegh. (). —
$Q-S$D676,:1$
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O%420#420&$V,686.6,$F-8=$@76$>+64-6>J$OH$42)C2%""$V,686.6,J$
@E$ 8, @+-4:. $ *E, -4:b$ 7+2)2C2+&)18:$V644;$ F-8=$ @76$ >+64-6>J$
7H$B2)%##%:":$V644;J$@E$T:.:G>-:b$7')20B2:"&$V.LC6$ F-8=$
8=,66$>+64-6>$@E$8,@+-4:.$*>-:$:7B$>-S$>+64-6>$@E$8,@+-4:.$*E,-4:b$
7')20B2:"21:":$Y7D.;$F-8=$ 8=,66$ >+64-6>$ @E$8,@+-4:.$ *E,-4:b$
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Fig. .
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F-8=$%/m20//m$?@@8>8,:+$>L++@,8$:7B$
V:G6>-:7$+@>86,-@,$+,@?:?-.8-6>$@E$0;/$
:,6$>=@F7$F-8=$?@.B$.-76>;$K=6$>-R6$@E$
8=6$8,-:7D.6$,6+,6>678>$7LC?6,$@E$D676,:$
+6,$E:C-.G$:7B$8=6$:48L:.$7LC?6,$->$D-AO
67$-7$+:,678=6>6>;$*6,-:.$+:,:>-8->C$->$
>=@F7$?G$B:,5$E -..$F-8=-7$8=6$8,-:7D.6>;$
]@@8$+:,:>-86>$:,6$B6+-486B$F-8=$F=-86$
E-..$F=6,6:>$7@7+:,:>-86>$:,6$>=@F7$-7$
D,6G;$K=6$8,@+=-4$7:8L,6>$@E$[48@576O
C:46:6$:7B$*+8:7B,:46:6$:,6$4L,,678.G$
L757@F7$?L8$:,6$>=@F7$:>$+:,:>-8-4$
:>>LC-7D$:$>-7D.6$@,-D-7$@E$+:,:>-8->C$
:7B$8=6-,$+@>-8-@7>$@7$8=6$8,66;

!^"
3-45,678$9$:.;$)$Classif ication of SantalalesTA X ON $!%$&'($)$*+,-.$'/0/1$!"#2!!#
9$:.;J$'//^(;$*$+@8678-:.$>G7:+@C@,+=G$E@,$8=6$4.:B6$->$:7->@O
4G8-4J$4G4.@4G8-4$>8@C:8:;$<E$:BB-8-@7:.$F@,5$EL,8=6,$>L++@,8>$
8=->$,6.:8-@7>=-+J$ ,64@D7-8-@7$ @E$@76$E:C-.GJ$Y,G8=,@+:.:46:6$
&F-8=$8F@$>L?E:C-.-6>(J$C:G$?6$PL>8-E-6B;
. Family Coulaceae Tiegh. (). —
$K=,66$[.B$_@,.B$
D676,:1$A28*&$V:-..;$F-8=$@76$>+64-6>J$AH$#C8*":$V:-..;$@E$8,@+-O
4:.$_;$*E,-4:b$J"%a8&)'"&$*L?.;$F-8=$@76$>+64-6>J$JH$48"&%#%G
:":$*L?.;J$@E$8,@+-4:.$*C6,-4:b$:7B$N+,&%2:'&+,5:$T:>8;J$F-8=$
@76$>+64-6>J$NH$&0#%'&+#&$T:>8;J$-7$F6>86,7$T:.:G>-:;
T6C?6,>$ @E$8=->$ E:C-.G$ :,6$ C6B-LC$ 8@$ .:,D6$8,66>$ F-8=$
:.86,7:86J$>-C+.6J$+68-@.:86J$6S>8-+L.:86$ .6:A6>;$K=6$E:C-.G$->$
C:,56B$ ?G$ >6A6,:.$ :7:8@C-4:.$ >G7:+@C@,+=-6>$ >L4=$ :>$.6:E$
B67B,-8-4$=:-,>J$.-D7-E-6B$ >64@7B:,G$F:..>$ @E$ 8=6$ 6+-B6,C->J$
.6:E$.:8-4-E6,>J$:7B$ 6+-B6,C:.$B,L>6>;$<7E.@,6>46746>$@44L,$:>$
>+-56O.-56$8=G,>6>;$K=6$?->6SL:.$E.@F6,>$:,6$B-+.@>86C@7@L>$8@$
+@.G>86C@7@L>$:7B$868,:O$@,$+678:C6,@L>;$K=6$>6+:.>$:,6$B6O
A6.@+6B$:7B$4@77:86$?L8$B@$7@8$E@,C$:7$:44,6>4678$4:.GS$L+@7$
E,L-8-7D;$K=6$D.:?,@L>$@,$+L?6>4678$+68:.>$C:G$?6$:+@+68:.@L>$
@,$>GC+68:.@L>$@7.G$:8$8=6$?:>6;$Q8:C67>$@44L,$-7$8F@$:7B$8=,66$
F=@,.>J$8=6$ 7LC?6,$6NL:.-7D$8=6$ 7LC?6,$@E$+68:.>$@,$=-D=6,$
&>8 :C - 7@B 6> $ >@ C68 -C 6> $ +, 6>6 78( ;$ *$ D .: 7B L. :, $ B-> 5$ - >$ : ?>6 78 ;$
K=6$@A:,G$->$=G+@DG7@L>$F-8=$8=,66$:7B$E@L,$ .@4L.6>;$ a,L-8>$
:,6$C6B-LC$8@$.:,D6$B,L+6>b$8=@>6$@E$A28*&$6:867$?G$=LC:7>;
K=->$A6,G$=@C@D676@L>$D,@L+$@E$8=,66$D676,:$F:>$,64@DO
7-R6B$:>$6:,.G$:>$0#%%$&K-6D=6CJ$0#%%:J?($:7B$?G$>L?>6NL678$
F@,56,>$&Q.6LC6,J$0%"!:b$Q8:LEE6,J$0%W0?($:7B$ F:>$4.:>>-E-6B$
-7$8,-?6$`@L.6:6;$V@8=$C@,+=@.@D-4:.$:7B$C@.64L.:,$:7:.G>6>$
&T:.c4@8$ 9$ :.;J$ '//^b$T:.c4@8$ 9$ 3-45,678J$'//#($=:B$ =-D=$
?@@8>8,:+$>L++@,8$E@,$8=6$ 4.:B6$,6+,6>678-7D$8=->$E:C-.G;$*.O
8=@LD=$8=6$4.:B6$->$4.6:,.G$C@7@+=G.68-4J$-8$6C6,D6>$:>$+:,8$@E$
:$+@.G8@CG$F-8=$8=6$76S8$8=,66$E:C-.-6>;
. Family Ximeniaceae Horan. (). —
$a@L,$ D676,:1$
A8)81")&$I;*;$V.:45$F-8=$@76$>+64-6>J$AH$'#.##%":$I;*;$V.:45$
@E$ *C:R@7-:7$ V,:R-.b$!28)&C2&$Q.6LC6,$F-8=$ @76$ >+64-6>J$
!H$+2%:"0"*":$Q.6LC 6,J$@E$V, : R-.$&* C :+: J$H: , :(b $J&*&%"&$`=L7$
9$Q;\;$ U66J$ F-8=$ @76$>+64-6>J$JH$2*#".#)&$ `=L7$ 9$Q;\;$U66J$
@E$_;$IL:7DS-$:7B$6:>86,7$nL77:7$ +,@A-746>$@E$`=-7:b$:7B$
`"0#%"&$U;$F-8=$867$>+64-6>$-7$8=6$[.B$:7B$36F$_@,.B$8,@+-4>;
T6C?6,>$@E$8=->$E:C-.G$:,6$>=,L?>$@,$>C:..$8@$.:,D6$8,66>$
F-8=$:.86,7:86J$>-C+.6J$+68-@.:86J$6S>8-+L.:86$.6:A6>;$`"0#%"&$
+@>>6>>$ :S-..:,G$ >+-76>$:7Bl@,$ ,:C:.$8=@,7>;$<7E .@,6>46746>$
:,6$EL7B:C678:..G$LC?6..:86$&,6BL46B$8@$>@.-8:,G$E.@F6,>$-7$
>@C6$>+64-6>$@E$`"0#%"&(;$K=6$?->6SL:.J$868,:C6,@L>$E.@F6,>$
=:A6$>6+:.>$8=:8$:,6$:,6$+:,8-:..G$@,$4@C+.686.G$4@77:86$?L8$B@$
7@8$?64@C6$:44,6>4678$L+@7$E,L-8-7D;$Q8:C67>$@44L,$-7$@76$@,$
8F@$F=@,.>$:7B$>8:C-7@B6>$:,6$:?>678;$K=6$@A:,G$->$>L+6,-@,$
F-8=$:$.@7D$4@7-4:.$>8G.6;$a,L-8>$ :,6$B,L+6>$F-8=$:$8=-7$+L.+G$
+6,-4:,+$:7B$:$4,L>8:46@L>$8@$F@@BG$67B@4:,+;
3@$C@.64L.:,$B:8:$F6,6$:A:-.:?.6$E@,$8=6$,:,6$!28)&C2&J$
=@F6A6,J$8=6$4.:B->8-4$:7:.G>->$?G$T:.c4@8$9$:.;$&'//^($+.:46B$
-8$-7$8=->$4.:B6;$K=6$E,L-8>$E,@C$J&*&%"&$+,@BL46$:7$6B-?.6$@-.J$
:.8=@LD=$-8$->$C@,6$4@CC@7.G$L>6B$:>$:$.L?,-4:78$E@,$C:4=-76,G$
&U66J$0%#/(;$Q-C-.:,.GJ$8=6$E:88G$E,L-8>$@E$A8)81")&$:,6$L>6B$8@$
C:56$>@:+$&Q.6LC6,J$0%#^(;
. Family Aptandraceae Miers (). —
$Y-D=8$D676,:1$
I%&+2*2:&$ &V.LC6($V.LC6$F-8=$ 0W$>+64-6>$-7$8=6$[.B$_@,.B$
8,@+-4>b$I1'&%C)&$T-6,>$F-8=$8=,66$>+64-6>$-7$8,@+-4:.$*C6,-4:$
:7B$@76$-7$*E,-4:b$A&',#C)&$T-6,>$F-8=$E-A6$>+64-6>$-7$Q@L8=$
*C6,-4:b$A,&8%2+,"'2%$V678=;$F-8=$8=,66$>+64-6>$-7$8,@+-4:.$
*C6,-4:b$M&)0&%C"&$H-6,,6$6S$V:-..;$F-8=$@76$>+64-6>J$MH$0#G
\2%4#%:":$H-6,,6$6S$V:-..;$-7$QY$*>-:$:7B$T:.:G>-:b$N%42\#&$
H-6,,6$F-8=$@76$>+64-6>J$NH$42)#$H-6,,6$-7$8,@+-4:.$_6>8$*E,-4:b$
:7B$-,&%#)2C":+8:$ `:A:4@$ F-8 =$ 8=,66$ >+6 4-6>$ 67B6C-4$ 8@$
T:B:D:>4:,;$*.>@J$:$C@7@8G+-4$D67L>$E,@C$ Z@7BL,:>J$M2%G
C8)2C#%C)2%$8)+#2*&'80$`;$ i..@:J$ 3-45,678J$_=-86E@@,B$ 9$
M;$\6..GJ$=:>$,64678.G$?667$B6>4,-?6B$&i..@:$9$:.;J$-7$+,6>>(;
*..$C6C?6,>$:,6$>=,L?>$:7B$8,66>$F-8=$:.86,7:86J$>-C+.6J$
+68-@.:86J$6S>8-+L.:86$.6:A6>;$M3*$>6NL6746$B:8:$,6+@,86B$:,6$
:A:-.:?.6$E@,$:..$D676,:$-7$8=6$E:C-.G$:7B$,6.:8-@7>=-+>$:,6$EL..G$
,6>@.A6B$&i..@:$9$:.;J$-7$+,6>>(;$T@.64L.:,$:7:.G>6>$D:A6$8F@$
4.:B6>$F-8=-7$8=6$E:C-.GJ$8=6$E-,>8$4@C+@>6B$@E$I% &+2*2:&J$A&G
',#C)&J$:7B$-,&%#)2C":+8:H$QG7:+@C@,+=-6>$E@,$8=6>6$D676,:$
-74.LB6$.-D7-E -6B$ DL:,B$ 46..>J$ +68:.>$ F-8=$ :+-4:.$ 8=-4567-7D>J$
+@,@>6$:78=6,$B6=->46746J$ +,@.@7D6B$:78=6,$4@77648-A6>J$:7B$
B-+.@+@,:86$+@..67;$V@8=$I%&+2*2:&$:7B$A&',#C)&$=:A6$:7$:4O
4,6>4678$B->5$8=:8$>L,,@L7B>$8=6$C:8L,6$B,L+6;$*.8=@LD=$@76$
F@L.B$:>>LC6$8=:8$-,&%#)2C":+8:J$D-A67$-8>$7:C6J$:.>@$=:>$:7$
:44,6>4678$B->5J$8=6$>-8L:8-@7$-7$8=->$D67L>$->$C@,6$4@C+.6S;$
K=6$B->5$8->>L6$8=:8$->$=@C@.@D@L>$F-8=$I%&+2*2:&$:7B$A&G
',#C)&$->$7@8$:44,6>4678;$ <7>86:BJ$ 8->>L6>$8=:8$:,6$7@8$+,6>678$
:8$E .@F6,-7D$?L8$ B6A6.@+$L+@7$E,L-8-7D$?68F667$ 8=6$ B->5$ :7B$
8=6$4:.GS$&8=6$6S8,:B->4:.$:7B$-7BLA-:.$4L+L.6>($6S+:7B$D,6:8.G$
6A678L:..G$E@,C-7D$-7E.:86B$E6>8@@7>$8=:8$>L,,@L7B$8=6$B,L+6;$
T6C?6,>$@E$8=6$>64@7B$4.:B6J$F-8=$I1'&%C)&J$A,&8%2+,"'2%J$
M&)0&%C"&J$M2%C8)2C#%C)2%$:7B$N%42\#&J$:..$=:A6$A:.A:86$
:78=6,$B6=->46746$:7B$:7$:44,6>4678$4:.GS;$K=6$D676,:$I1'&%G
C)&J$M&)0&%C"&$: 7B$N%42\#&$:.>@$>=:,6 $8=6$4=:, : 486 ,$@E$EL>6 B$
>8:C-7:.$E-.:C678>J$:$D.:7BL.:,$B->5$?68F667$8=6$>8:C67>$:7B$
+68:.>J$:7B$:$4@74:A6$:+@4@.+-LC$@7$8=6$+@..67;$*..$C6C?6,>$
@E$8=6$E:C-.G$=:A6$:$?:>:..G$?-.@4L.:,$@A:, GJ$:.86,7:86$-786,A:>O
4L.:,$+-8>J$:7B$?,6A-:S-:.$+@..67$D,:-7>;$<8$->$@E$-786,6>8$8=:8$8=6$
867B674G$8@F:,B>$ :44,6>46746$ -7$ *+8:7B,:46:6$@44L,>$ -7$ :..$
C6C?6,>$ ?L8$ 8=6$ :EE6486B$ 8->>L6>$A:,G;$K=6$ A:,-:8-@7$ :C@7D$
8:S:$-7$8=6$C@,+=@D6768-4$6EE648J$E,@C$8=6$-776,$&B->5(J$8@$8=6$
C-BB.6$ &-7BLA-:.$ 4L+L.6($8@$ 8=6$ @L86,$&4:.GS($+@,8-@7>$ @E$8=6$
,646+8:4.6$C:G$ ,6+,6>678$ :$D,:B-678$,6>+@7>6$ 8@$ :$8,:7>4,-+O
8-@7$E:48@,;$i7.-56$8G+-4:.$E .@,:.$=@C6@8-4$CL8:78>J$=@F6A6,J$
8=->$+,@46>>$B@6>$7@8$,6>L.8$-7$8=6$.@>>$@E$E.@,:.$@,D:7>$&@,$8=6-,$
8,:7>E@,C:8-@7$-78@$:7@8=6,($?L8$-7$8=6$6S+:7>-@7$@E$6S->8-7D$
@,$7@A6.$@,D:7>;
. Family Olacaceae R. Br. (), emend. —
$K=,66$D676,:1$
!8*&+"&$Q.6LC6,$F-8=$0"$>+64-6>$ @E$Q@L8=$*C6,-4:b$N*&_$U;$
F-8=$4:;$^/$>+64-6>$-7$[.B$_@,.B$8,@+-4>$:7B$*E,-4:b$:7B$-'5G
+,21#'&*80$V678=;$F-8=$8F@$>+64-6>$-7$8,@+-4:.$Q@L8=$*C6,-4:$
:7B$8F@$-7$8,@+-4:.$F6>86,7$:7B$4678,:.$*E,-4:;
T6C?6,>$ @E$ [.:4:46:6$>;>8,;$ :,6$ 8,66>J$ >=,L?>$@,$,:,6.G$
.-:7:>$F-8=$:.86,7:86J$>-C+.6J$+68-@.:86J$D.:?,@L>J$6S>8-+L.:86$
.6:A6>;$K=6$6.@7D:86$E.@F6,>$&@A:.$-7$?LB($:,6$+678:C6,@L>$@,$
>@C68-C6>$868,:O$@,$=6S:C6,@L>;$*$4:.GS$@,$4:.G4L.L>$->$+,6>O
678$&-7$N*&_$:7B$!8*&+"&($F=-4=$?64@C6>$:44,6>4678J$8-D=8.G$
>L,,@L7B-7D$8=6$E,L-8;$K=6$:7B,@64-LC$4@C+,->6>$8F@$F=@,.>$
8=:8$@E867$-74.LB6$>8:C-7@B6>;$*$D.:7BL.:,$B->5$->$:?>678;$K=6$
@A:,G$ ->$ >L+6,-@,$@,$=:.EO-7E6,-@,J$4@C+@>6B$@E$8=,66$ 4:,+6.>$
?L8$F-8=$@76$.@4L.6;$a,L-8>$ :,6$ B,L+6>$ @,$ +>6LB@OB,L+6>J$ 8=6$

!^^
TA XO N$!%$&'($)$*+,-.$'/0/1$!"#2!!#3-45,678$9$:.;$)$Classif ication of Santalales
.:886,$ 86,C$ L>6B$?G$Q.6LC6,$&0%#^($8@$:44@L78$ E@,$ 8:S:$F-8=$
4@74,6>4678$8->>L6>;
K=->$ E:C-.GJ$E-,>8$ ,64@D7-R6B$ ?G$ K-6D=6C$ &0#%W(J$>=:,6>$
>6A6,:.$:7:8@C-4:.$&6;D;J$>-.-4:$?@B-6>$-7$F@@B$,:G>$46..>(J$C@,O
+=@.@D-4:.J$:7B$+:.G7@.@D-4:.$>G7:+@C@,+=-6>$&T:.c4@8$9$:.;J$
'//^(;$T@.64L.:,$B:8:$.-75$8=6$D67L>$!8*&+"&$8@$* E , -4: 7$C6 CO
?6,>$@E$N*&_$>648;$P)"&%C ) & #$Y7D.;$&T:.c4@8$9$3-45,678J$'//#(;$
<7B66BJ$ :$ C@.64L.:,$+=G.@D6768-4$-7A6>8-D:8-@7$ 8=:8$-74.LB6>$
,@?L>8$>:C+.-7D$@E$?@8=$N*&_$:7B$!8*&+"&$->$,6NL-,6B$8@$B686,O
C-76$F=68=6,$8=6>6$8F@$D676,:$:,6$B->8-748$@,$7@8;
. Family Octoknemaceae Soler (). —
$T@7@D676,-4J$
F-8=$N+'2\%#0&$H-6,,6$ =:A-7D$ 4:;$ >6A67$ >+64-6>$@E$8,@+-4:.$
*E,-4:;
N+'2\%#0&$-74.LB6>$B-@64-@L>$8,66>$ :7B$>=,L?>$F-8=$:.O
86,7:86J$>-C+.6J$+68-@.:86J$6S>8-+L.:86$ .6:A6>$ F-8=$>86..:86$+LO
?6>46746;$K=6$-7E.@,6>46746>$:,6$E:>4-4L.:86$:7B$6C6,D6$E,@C$
67B@D67@L>$?LB>$?6.@F$8=6$?:,5;$T:.6$E .@F6,>$:,6$+678:CO
6,@L>$F-8=$ >C:..$ 4:.GS$8668=$:.86,7:8-7D$ ?68F667$8=6$A:.A:86$
+68:.>;$*$ .@?6B$D.:7BL.:,$B->5$>L,,@L7B>$8=6$+->8-..@B6;$K=6$
+678:C6,@L>$E6C:.6$E .@F6,>$?6:,$ :$>8:C-7@B6$@++@>-86$6:4=$
+68:.$:7B$=:A6$:$CL.8-.@?6B$>8-DC:$:8@+$8=6$>8G.6;$K=6$-7E6,-@,$
@A:,G$ ->$ "O.@4L.:,$?6.@F$?L8$ ?64@C6>$L7-.@4L.:,$:?@A6;$K=6$
E,L-8$->$:$B,L+6$4,@F76B$?G$8=6$+6,>->8678$+6,-:78=;$K=6$6S86,-@,$
@E$8=6$>66B$->$B66+.G$,LC-7:86O.@?6B;
K=->$67-DC:8-4$8:S@7$=:>$4=:..67D6B$4.:>>-E-4:8-@7$E@,$@A6,$
0// $G6: ,>;$K =6$D67L >$F: >$+.:46 B$-7$-8>$@F7$E:C-.G$?G$K-6D =6C$
&0%/ !($: 7B$= : >$>- 746$?667 $:..-6B$F -8=$[.: 4:46 : 6$?G$C@ >8$F@, 5O
6,>;$U@L->$9$Uc@7:,B$&0%^#($,64@D7-R6B$[48@576C:46:6$:>$:$
E:C-.GJ$?L8$-74.LB6B$F-8=-7$-8$N\28B&\&$&=6,6$-7$`6,A:786>-:O
46:6(;$T@,+=@.@D-4:.$E6:8L,6>$:,6$6-8=6,$L7L>L:.$E@,$Q:78:.:.6>J$
>L4=$:>$6S+:7B6B$>8-DC:8-4$6S4,6>46746>$-7$8=6$E6C:.6$E.@F6,>J$
@,$+,@A-B6$4@7E.-48-7D$.-75>$8@$A:,-@L>$D,@L+>;$a@,$6S:C+.6J$8=6$
E.@,:.$>86..:86$8,-4=@C6>$:7B$F@@B$:7:8@CG$.-75$8@$`@L.:46:6$
&T-.B?,6:BJ$0%"!b$]66BJ$0%!!(;$U6:E$:7B$F@@B$:7:8@CG$&V::>J$
0%#'b$A:7$B67$[6A6,J$0%#^($-7B-4:86$8=->$8:S@7$->$->@.:86B$E,@C$
@8=6,$[.:4:46:6;$H@..67$L.8,:>8,L48L,6$>LDD6>86B$:$,6.:8-@7>=-+$
F-8=$[+-.-:46:6$&U@?,6:LO`:..67J$0%#'(J$:$C@,6$B6,-A6B$+@>-O
8-@7$F-8=-7$8=6$@,B6,$8=:8$E@..@F>$Q4=L.8R6OT@86.$&0%W^($F=@$
C:B6$-8$:$8,-?6$-7$>L?E:C-.G$Q4=@6+E-@-B6:6;$K=6$,6BL48-@7$@,$
.@>>$ @E$ >6+:.>$ -7$ >@C6$N\'2\%#0&$>+64-6>$ ->$ -7$:44@,B$ F-8=$
8,67B>$>667$-7$C@,6$B6,-A6B$C6C?6,>$@E$Q:78:.:.6>;$K=6$+@>-O
8-@7$@E$N+'2\%#0&$@7$8=6$C@.64L.:,$8,66$&T:.c4@8$9$3-45,678J$
'//#($:>$>->86,$&?L8$F-8=$.@F$>L++@,8($8@$:$4.:B6$4@C+@>6B$@E$
Q4=@6+E-:46:6J$T->@B67B,:46:6J$U@,:78=:46:6$:7B$[+-.-:46:6$
:.>@$>L++@,8>$8=->$4@746+8;$H:,:>-8->C$=:>$7@8$?667$B@4LC6786B$
E@,$ N+'2\%#0&J$ ?L8$ -8>$ +=G.@D6768-4$ +@>-8-@7$ +@-78>$8@F:,B>$
8=->$8,@+=-4$4@7B-8-@7;
. Family Schoepfiaceae Blume (). —
$K=,66$D676,:1$
I)b2%&$`@CC;$6S$`:A;$F-8=$867$>+64-6>$@E$8,@+-4:.$:7B$86C+6,O
:86$Q@L8=$*C6,-4:b$W8"%+,&0&*"80$T@.-7:$F-8=$4:;$'/$>+64-6>$
@E$*7B6:7$Q@L8=$*C6,-4:b$:7B$7+,2#1."&$Q4=,6?;$F-8=$4:;$'!$
>+64-6>$-7$8,@+-4:.$*C6,-4:J$*E,-4:J$:7B$*>-:;
Q4=@6+E-:46:6$-74.LB6$+.:78>$F-8=$B-A6,>6$=:?-8>1$,@@8O+:,O
:>-8-4$8,66>J$>L?>=,L?>$F-8=$SG.@+@B-:$&I)b2%&$>648;$`5*&)b2%&$
Q5@88>?6,D($:7B$=6,?:46@L>$+6,677-:.>$F-8=$:.86, 7:86J$>-C+.6J$
6S>8-+L.:86$.6:A6>;$K=6$?->6SL:.$E.@F6,>$:,6$?@,76$-7$:S-..:,G$
:7B$86,C-7:.$-7E.@,6>46746>$@E$A:,-@L>$8G+6>;$K=6$E.@,:.$?,:48>$
:7B$?,:486@.6>$ :,6$ +6,>->8678J$>6+:,:86$@,$EL>6B$ -78@$:$ 4L+$ -7$
W8"%+,&0&*"80$:7B$7+,2#1."&;$K=6$4:.GS$->$,6+,6>6786B$?G$:$
4:.G4L.L>$-7$7+,2#1."&;$Z:-,>$@44L,$@++@>-86$8=6$>8:C67>$@7$8=6$
^$@,$!$+68:.>$@E$I)b2%&$:7B$7+,2#1."&;$K=6$@A:,G$->$-7E6,-@,$F-8=$
'$@,$"$.@4L.6>$:8$8=6$?:>6$:7B$L7-.@4L.:,$:?@A6;$K=6$C:8L,6$E,L-8$
-7$7+,2#1."&$->$:$B, L+6 $F=6 ,6:> $-7$I)b2%&$:7B $W8"%+,&0&*"80$
-8$->$:$7L8.68J$4,@F76B$?G$+6,>->8678$+6,-:78=$+:,8>;$K=6$?,:48>$
8=:8$674.@>6$8=6$ E,L-8$ :,6$:44,6>4678$F-8=$:$>4.6,@8-4$ .:G6,$-7$
I)b2%&$:7B$W8"%+,&0&*"80;
K=6$:>>@4-:8-@7$@E$I)b2%&$:7B$W8"%+,&0&*"80$=:>$76A6,$
?667$B->+L86BJ$=@F6A6,J$8=6-,$:EE-7-8G$F-8=$7+,2#1."&J$:>$@,-D-O
7:..G$7@86B$?G$K-6D=6C$&0#%W(J$F:>$@7.G$,64678.G$4@7E-,C6B$
L>-7D$C@.64L.:,$+=G.@D6768-4$C68=@B>$&M6,$9$3-45,678J$'//#(;$
K=->$,6.:8-@7>=-+$->$>L++@,86B$?G$8=6$:?@A6$C@,+=@.@D-4:.$E6:O
8L,6>;
. Family Misodendraceae J. Agardh (). —
$T@7@D6O
76,-4J$F-8=$J":2C#%C)80$V:75>$6S$M`;$=:A-7D$6-D=8$>+64-6>$
@E$:6,-:.$=6C-+:,:>-8-4$>=,L?>$67B6C-4$8@$86C+6,:86$E@,6>8>$@E$
Q@L8=$*C6,-4:J$C:-7.G$>@L8=$@E$""$B6D,66>;
V,:74=-7D$ -7$J":2C#%C)80$ ->$ >GC+@B-:.$ :7B$ 8=6$.6:A6>$
:,6$:.86,7:86;$H.:78>$:,6$B-@64-@L>$&,:,6.G$C@7@64-@L>$@,$F-8=$
?->6SL:.$E.@F6,>(;$K=6$-7E.@,6>46746>$:,6$@E$>+-56>$@,$,:46C6>$
8=:8$?6:,$,6BL46B$E.@F6,>$>L?867B6B$?G$?,:48>;$K=6$>8:C-7:86$
E.@F6,>$ .:45$:$+6,-:78=$ :7B$=:A6$:$4678,:.$7648:,-E6,@L>$B->5$
>L,,@L7B6B$ ?G$ 8F@$ @,$ 8=,66$ >8:C67>;$K=6$ 4:,+6..:86$ E .@F6,>$
:,6$6+-DG7@L>$&7@8$=G+@DG7@L>J$>66$d-B:.O]L>>6..$9$3-45,678J$
'//X($:7B$8=6$>8:C-7@B6>$&@,$>8:C67>$-7$?->6SL:.$E.@F6,>($:,6$
EL>6B$8@$8=6$@A:,G$@7.G$:8$8=6$?:>6$&6+-=G+@DG7@L>(;$K=6$>8:O
C-7@B6>$6C6,D6$E,@C$D,@@A6>$?68F667$8=6$+6,-:78=$C6C?6,>;$
H6,-:78=$.@?6>$&+68:.>($:,6$,64@D7-R:?.6$:8$8=6$:+6S$@E$8=6$@A:,G$
-7$>@C6$>+64-6>;$*$7648:,-E6,@L>$B->5$@44L,>$?68F667$8=6$+68O
:.>$:7B$8=6$>8G.6$F-8=$8=,66$>8-DC:>;$K=6$C:8L,6$:4=676$?6:,>$
8=,66$:44,6>4678$E6:8=6,G$>8:C-7@B6>$8=:8$566+$8=6$E,L-8$:.@E8$
:7B$:.>@$:-B$-7$:88:4=C678$8@$8=6$=@>8$?,:74=;
*..$,64678$CL.8-D676$C@.64L.:,$:7:.G>6>$&M6,$9$3-45,678J$
'//#b$T:.c4@8$9$3-45,678J$ '//#b$d-B:.O]L>>6..$9$3-45,678J$
'//#?($+.:46$T->@B67B,:46:6$:>$>->86,$8@$Q4=@6+E-:46:6$:7B$
8=:8$4.:B6$>->86,$8@$U@,:78=:46:6;$*C@7D$8=6$E-A6$-7B6+67B678$
6A@.L8-@7>$@E$:6,-:.$+:,:>-8->CJ$C@.64L.:,$B:8-7D$-7B-4:86$J":2G
C#%C)80$E-,>8$:4=-6A6B$8=->$=:?-8$4:;$#/$T:$&d-B:.O]L>>6..$9$
3-45,678J$'//#?(;
. Family Loranthaceae Juss. (). —
$Q6A678GO8=,66$
D676,:$C:-7.G$@E$8=6$[.B$:7B$36F$_@,.B$8,@+-4>$F-8=$ >@C6$
D676,:$@44L,,-7D$-7$86C+6,:86$,6D-@7>;$K=6$>L?E:C-.-:.$7:C6$
U@,:78=@-B6:6$Y:8@7$&0#"W($F:>$L>6B$?G$Y7D.6,$9$\,:L>6$
&0%"!($E@,$8=6$8:S@7$6NL-A:.678$8@$F=:8$->$=6,6$4@7>-B6,6B$E:CO
-.G$U@,:78=:46:6;$K=6$@8=6,$>L?E:C-.GJ$d->4@-B6:6J$->$=6,6$,64O
@D7-R6B$:>$ :$B->8-748$E:C-.G$&>66$B->4L>>-@7$ L7B6,$d->4:46:6$
?6.@F(;
Z6C-+:,:>-8-4$@7$8=6$,@@8>$:7Bl@,$?,:74=6>$@E$=@>8$+.:78>$
?G$ C6:7>$ @E$ +,-C :,G$ @,$ >64@7B:, G$ =:L>8@, -:$ :7B$ F-8=$ @,$
F-8=@L8$ 6+-4@,8-4:.$ ,L776,>;$K=6$ .6:A6>$ :,6$C@>8.G$@++@>-86$
&C@,6$,:,6.G$: .8 6,7:86$@,$F=@,.6B(J$>-C+.6J$4L , A - 76, A6B$@,$+67O
7-76,A6B;$<7E .@,6>46746>$:,6$ 86,C-7:.$:7B$.:86,:.J$ -7$A:,-@L>$
E@,C>$ -74.LB-7D$ >+-56>J$ ,:46C6>J$LC?6.>J$:7B$4:+-8 L.:J$ ?L8$
8=6$?:>-4$L7-8$->$@E867$:$B-4=:>-LC;$a.@F6,>$:,6$?->6SL:.$@,$
L7->6SL:.J$,:B-:..G$@,$?-.:86,:..G$>GCC68,-4:.J$^2W&2#(OC6,@L>J$

!^!
3-45,678$9$:.;$)$Classif ication of SantalalesTA X ON $!%$&'($)$*+,-.$'/0/1$!"#2!!#
F-8=$:$4:.G4L.L>$&4:.GS$,6BL46B$8@$:$,-CJ$:$8L?6J$@,$>=@,8$8668=(J$
:7B$:$D:C@+68:.@L>$@,$4=@,-+68:.@L>$4@,@..:J$F-8=$A:.A:86$+68O
:.>;$K=6$>8:C67>$:,6$:>$C:7G$:>$8=6$+68:.>$:7B$6+-+68:.@L>J$8=6$
:78=6,>$:,6$'O$@,$^O.@4L.:,J$?:>-E-S6B$@,$B@,>-E-S6B;$K=6$@A:,G$
->$-7E6,-@,J$L7-.@4L.:,$&@,$-8>$ .@4L.6>$>@C68-C6>$=:A6$@?>4L,6$
.@?6>$:8$8=6$?:>6(J$:7B$.:45>$B-EE6,678-:86B$@AL.6>$&8=6$6C?,G@$
>:4>$E@,C$F-8=-7$:$ C:C6.@7(;$K=6$E,L-8$->$ :$ ?6,,G$ &6S46+8$-7$
S85':"&($F-8=$:$>-7D.6$A->4-B$>66B;
K=,66$ D676,:J$S85':"&J$I'\"%:2%"&$:7B$ L&"&C#%C)2%$:,6$
>@.6.G$@,$+,-C:,-.G$,@@8$+:,:>-86>$F=6,6:>$8=6$C:P@,-8G$@E$8=6$
,6C:-7-7D$D676,:$:,6$>86C$+:,:>-86>$&C->8.68@6>(;$]@@8$:7B$>86C$
+:,:>-8->C$&8=6$:C+=-+=:D@L>$4@7B-8-@7J$>66$d-B:.O]L>>6..$9$
3-45,678J$'//#:J?($4:7$ ?6$E@L7B$-7$A:,-@L>$ >+64-6>$@E$P)"12G
C&%',8:$:7B$M#*"_&%',#)&;$K=6$E:C-.G$ ->$ >8,@7D.G$>L++@,86B$
:>$C@7@+=G.68-4$:>$->$8=6$ >->86,$,6.:8-@7>=-+$?68F667$S85':"&$
:7B$8=6$,6C:-7-7D$D676,:;
U@,:78=:46:6$:,6$=6,6$B-A-B6B$-78@$8,-?6>J$:7B$-7$>@C6$
4:>6>$>L?8,-?6>$&a-D;$'(;$*.8=@LD=$8=6$C@.64L.:,$8,66$ &d-B:.O
]L>>6..$ 9$ 3-45,678 J$ '//#:($=: >$ =-6,:,4=-4:.$ >8 ,L48L ,6$ 8=:8$
4@L.B$?6$L>6B$ 8@$+,@+@>6$ :BB-8-@7:.$ ,:75>$ &-;6;J$D,@L+-7D$ @E$
K,-?6>(J$-7$>@C6$4:>6>$,6.:8-@7>=-+>$:C@7D$8=6$4.:B6>$:,6$7@8$
F6..$>L++@,86BJ$=6746$8=6>6$F-..$7@8$?6$+,@+@>6B$=6,6;$K=L>J$
8=6$E@..@F-7D$-7E,:E:C-.-:.$4.:>>-E-4:8-@7$8:56>$-78@$:44@L78$8=6$
C@.64L.:,$8,66$8@+@.@DGJ$4.:B6$>L++@,8J$C@,+=@.@DGJ$:7B$?:>6$
4=,@C@>@C6$7LC?6,;$I-A67$8=6$8@+@.@DG$@E$8=6$C@.64L.:,$8,66$
&a-D;$'(J$-8$:++6:,>$8=:8$E@L,$C:P@,$:76L+.@-B$,6BL48-@7>$E,@C$%$j$
0'$@44L,,6B$8=,@LD=@L8$8=6$6A@.L8-@7:,G$=->8@,G$@E$8=6$E:C-.G1$
E-,>8$-7$@"4&)"&$8@$%$j$0/J$>64@7B$-7$>L?8,-?6$H>-88:4:78=-7:6$8@$
%$j$#J$8=-,B$-7$>L?8,-?6$<.6@>8G.-7:6$8@$%$j$00J$:7B$E@L,8=$-7$8=6$
4.:B6$4@C+,->-7D$ >L?8,-?6>$ U@,:78=-7:6J$ *CG6C-7:6J$ Q4L,O
,L.-7:6J$M67B,@+=8=@-7:6J$YC6.-:78=-7:6J$:7B$K:+-7:78=-7:6J$
:..$%$j$%;
Fig. .
$M-:D,:C:8-4$+=G.@D6768-4$8,66$>=@F-7D$,6.:8-@7>=-+>$:C@7D$8=6$8 ,-?6>$:7B$>L?8,-?6>$@E$U@,:78=:46:6;$K=->$8,66$->$?:>6B$+,-C:,-.G$L+@7$
8=6$C@.64L.:,$+=G.@D6768-4$-7A6>8-D:8-@7$?G$d-B:.O]L>>6..$9$3-45,678$&'//#:(;$3@B6>$,6>@.A6B$F-8=$0//m$?@@8>8,:+$>L++@,8$:7Bl@,$V:G6>-:7$
+@>86,-@,$+,@?:?-.8-6>$@E$0;/$:,6$>=@F7$F-8=$?@.B$.-76>J$#/m2%%m$F-8=$B:>=6B$.-76>J$X/m2X%m$@+67$.-76>J$:7B$oX/m$F-8=$8=-7$.-76>;$*BB-8-@7:.$
F6..O>L++@,86B$8@+@.@D-6>$C:G$6S->8$F-8=-7$8=6$.:,D6,$4.:B6>$?L8$:,6$7@8$>=@F7$&>66$d-B:.O]L>>6..$9$3-45,678J$'//#:(;$I676,:$7@8$>:C+.6B$-7$8=6$
C@.64L.:,$ >8LB-6>$&674.@>6B$ F-8=-7$+:,678=6>6>($:,6$+.:46B$-7$4.:B6>$?:>6B$L+@7$C@,+=@.@D-4:.$6A-B6746;$K=6$E@L,$:76L+.@-B$,6BL48-@7>$E,@C$
%$j$0'$:,6$-7B-4:86B$F-8=$:,,@F>$&-7$>L?8,-?6$U-D:,-7:6$%$j$0/$:++.-6>$@7.G$8@$@"4&)"&b$P) ":' # )"_$,6C:-7>$%$j$0'(;

!^W
TA XO N$!%$&'($)$*+,-.$'/0/1$!"#2!!#3-45,678$9$:.;$)$Classif ication of Santalales
.. Tribe Nuytsieae Tiegh. —
$K=->$8,-?6J$,64@D7-R6B$?G$
M:7>6,$&0%""(J$-74.LB6>$@7.G$S85':"&$];$V,;$F-8=$@76$>+64-6>J$
SH$.*2)"B8%C&$&U:?-..;($];$V,;J$@E$F6>86,7$*L>8,:.-:;
_-8=-7$U@,:78=:46:6J$S85':"&$=:>$7LC6,@L>$L7-NL6$E6:O
8L,6>$>L4=$ :>$ -8>$E,L-8$ &:$B,G$ 8=,66OF-7D6BJ$F-7BOB->+6,>6B$
>:C:,:(J$:7B$L7-NL6$=:L>8@,-:.$?6=:A-@,$:7B$:7:8@CG$&V6G6,$
9$:.;J$ 0%#%b$`:..:B-76$9$H:86J$'///(;$H.:78>$ :,6$C@7@64-@L>$
F-8=$:$?->6SL:.$E.@F6,$4678,:.$-7$8=6$B-4=:>-LC$:7B$C:.6$.:8O
6,:.$E .@F6,>;$a.@F6,>$?6:,$:$4:.G4L.L>$&F-8=$A:>4L.:,$>8,:7B>($
:7B$>-S$D:C@+68:.@L>$+68:.>$6:4=$F-8=$:$>8:C67;$*8$8=6$?:>6$
@E$ 8=6$ -7E6,-@,$ @A:,G$ :,6$ 8=,66$ >6C-.@4L.6>$ ?L8$ 8=6$ @A:,G$ ->$
L7-.@4L.:,$:?@A6;
.. Tribe Gaiadendreae Tiegh. —
$K=->$ 8,-?6$ -74.LB6>$
I'\"%:2%"&$a; $ TL 6. .; $ F- 8= $ @7 6$ >+ 64 -6 >J $ IH$*"48:')"%&$ &U-7B.;($
a;$TL6..;J$67B6C-4$8@$8=6$V.L6$T8>;$@E$36F$Q@L8=$_:.6>J$*L>O
8,:.-:J$:7B$L&"&C#%C)2%$I;$M @7$F -8=$@76 $>+64-6 >J$LH$18%+'&'80$
&]L-R$9$H:A@7($I;$M@7J$@E$`678,:.$:7B$Q@L8=$*C6,-4:;
I'\"%:2%"&$->$:$0$C$8:..$>=,L?$F-8=$@++@>-86J$D.:?,@L>J$
.:746@.:86$ .6:A6>;$ <8$ ?6:,>$ :7$ :S-..:, G$ ,:46C6$ @E$WOC6,@L>$
D:C@+68:.@L>$E .@F6,>$8=:8$8L,7$E,@C$G6..@F$8@$@,:7D6$:>$8=6G$
C:8L,6;$L&"&C#%C)2%$->$:$>=,L?$@,$8,66$L+$8@$0^$C$8:..J$6S->8O
-7D$6-8=6,$:>$:$86,,6>8,-:.$,@@8$+:,:>-86$@,$:>$:7$6+-+=G86$&\L-P8J$
0%W"(;$<8$=:>$@++ @>-8 6$8@$>L?@++@>-8 6$.6:A6>$:7B$?6 :,>$,:46C @>6$
@,$+:7-4L.:86$-7E .@,6>46746>$ 4@78:-7-7D$B-4=:>-:J$ F-8=$6:4=$
8,-:B$>L?867B6B$?G$8F@$>C:..6,$:7B$@76$.:,D6,$?,:48;$V@8=$D67O
6,:$=:A6$A:>4L.:,$>8,:7B>$-7$8=6$4:.G4L.L>J$B,L+:46@L>$E,L-8>J$
EL,,@F6B$67B@>+6, CJ$ :7B$ A6,>:8-.6J$ B@,>-E -S6B$ :78=6,>;$ K=6$
-7E .@,6>46746$ @E$I'\"%:2%"&$->$4@C+@>6B$@E$C@7:B>$F=6,6:>$
L&"&C#%C)2%$=:>$8,-:B>J$?L8$@8=6,F->6$8=6$E.@,:.$C@,+=@.@DG$-7$
8=6>6$D676,:$->$A6,G$>-C-.:,$:7B$C:84=6>$F6..$8=6$=G+@8=68-4:.$
:746>8,:.$8G+6$8=:8$C:G$=:A6$=:B$:7$@+67J$G6..@FJ$4=@,-+68:.O
@L>J$678@C@+=-.@L>$E.@F6,$&d-B:.O]L>>6..$9$3-45,678J$'//#:(;
Y7D.6,$&0#%X:($:7B$M:7>6,$&0%""($4:..6B$8=->$D,@L+$>L?8,-?6$
I:-:B67B,-7:6$F=6,6:>$K-6D=6C$&0#%!:($ ,64@D7-R6B$ -8$:>$:$
8,-?6;$T@.64L.:,$B:8:$B@$7@8$4@7>->8678.G$,6>@.A6$8=6>6$8F@$D67O
6,:$:>$:$4.:B6$?L8$:>$:$>L446>>-A6$D,:B6;$*BB-8-@7:.$C@.64L.:,$
F@,5$->$766B6B$8@$4@7E-,C$8=6-,$C@7@+=G.G;$K=6G$:,6$+.:46B$-7$
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.. Tribe Elytrantheae Engl. —
$K=->$8,-?6J$,64@D7-R6B$
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8=6$H=-.-++-76>J$T@.L44:>$:7B$36F$IL-76:b$!#+&":%"%&$K-6D=;$
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.. Tribe Psittacantheae Horan. —
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... Subtribe Tupeinae Nickrent & Vidal-Russell, subtr.
nov. —
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Q4=.8B.;J$@E$36F$p6:.:7B;

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... Subtribe Notantherinae Nickrent & Vidal-Russell,
subtr. nov. —
$KG+61$S2'&%',#)&$&M`;($I;$ M@7;$ KF@$D676,:1$
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... Subtribe Ligarinae Nickrent & Vidal-Russell,
subtr. nov. —
$KG+61$@"4&)"&$K-6D =;$KF@ $D676,:J $@"4&)"&$K-6D=;$
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... Subtribe Psittacanthinae Engl. —
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8=:8$ ->$ +6,>->8678$ @7$ 8=6$E,L-8J$:7B$ >+=6,-4:.$&7@8$8,-:7DL.:,($
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->$,6C:,5:?.6$-7$8=:8$-8$?6D-7>$.-E6$:>$:7$6+-+=G86$?L8$.:86,$-8>$
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8,-:B>J$&:.>@$B-:B>$-7$I#'&%',8:$: 7B$-:"''&+&%',8:($:7B$8=6>6$4:7$
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:$4L+L.6$&-,',")8:&J$ -:"''&+&%',8:J$ 684;(;$K=6$:48-7@C@,+=-4$
E.@F6,>$:,6$?->6SL:.$@,$L7->6SL:.J$:7B$-7$8=6$.:886,$?@8=$>8:C-O
7@B6>$:7B$+->8-..@B6>$C:G$?6$+,6>678;$<7$>C:..OE.@F6,6B$D676,:$
8=6$+68:.>$:,6$E,66$@,$@7.G$>.-D=8.G$EL>6B$:8$8=6$?:>6$F=6,6:>$-7$
8=6$8F@$.:,D6OE.@F6,6B$D676,:$&I#'&%',8:$:7B$-:"''&+&%',8:(J$
:$.@7D$4@,@..:$8L?6$->$>667;$Q8:C67>$:,6$L>L:..G$B-C@,+=-4J$@4O
4L,,-7D$:8$B-EE6,678$=6-D=8>$F-8=-7$8=6$E.@F6,$?LB;$<7$-,',")8:&J$
+,6>>L,6$E,@C$>=@,8$>8:C67$:78=6,>$4:L>6>$-7B678:8-@7>$-7$8=6$
E-.:C678>$@E$8=6$.@7D6,$>8:C67>;$*78=6,>$4:7$?6$6-8=6,$?:>-E-S6B$
@,$B@,>-E-S6B$:7B$-7$>@C6$D676,:$8=6$4@77648-A6$->$:+-4L.:86;$
K=6$>8G.6$->$4@78@,86B$-7$A*&C2+2*#&$:7B$7')8',&%',8:;
T@.64L.:,$+=G.@D67-6>$:7B$+:.G7@.@D-4:.$6A-B6746$&a6L6,$9$
\L-P8J$0%#!($,6A6:.$8=6$4.@>6$,6.:8-@7>=-+>$:C@7D$8=6$>6A67$>C:..O
E.@F6,6B$D676,:$&A*&C2+2*#&J$!#%C)21#02%J$9_2+&+'8:J$N)5+G
'&%',8:J$N) 5 +'"% &J$-&%& 0&%' ,8:J$-,',")8:& J$7')8',&%',8:(J$8=L>$
EL8L,6$F@,5$->$766B6B$8@$C@,6$4.6:,.G$B6E-76$D676,-4$?@L7B:,-6>;

!^#
TA XO N$!%$&'($)$*+,-.$'/0/1$!"#2!!#3-45,678$9$:.;$)$Classif ication of Santalales
.. Tribe Lorantheae Rchb. —
$K=->$8,-?:.$ 7:C6$F:>$
:.>@$L>6B$?G$Y7D.6,$9$\,:L>6$&0%"!(;$K=->$->$8=6$.:,D6>8$8,-?6$
-7$U@,:78=:46:6$:7B$4@78:-7>$>6A67$>L?8,-?6>$:7B$^/$D676,:;$
Y7D.6,s>$>L?8,-?6$U@,:78=-7:6$-74.LB6B$>6A67$D676,:J$?L8$@76$
@E$ 8=6>6$ F:>$ @2)&%',8:$ 8=:8J$:44@,B-7D$8@$C@B6,7$4@746+8>J$
674@C+:>>6>$"X$D676,:;$QL?8,-?6$<.6@>8G.-7:6$=:>$:$?:>6$4=,@O
C@>@C6$7LC?6,$ @E$ _$j$00$:7B$8=->$4.:B6$->$>->86,$8@$:$4.:B6$
4@78:-7-7D$8=6$,6C:-7-7D$>-S$>L?8,-?6>$&"#$D676,:($8=:8$:..$=:A6$
_$j$%;$K=->$,6+,6>678>$8=6$E@L,8=$:76L+.@-B$,6BL48-@7$-7$U@O
,:78=:46:6J$:7B$@76$8=:8$:EE6486B$8=6$.:,D6>8$7LC?6,$@E$D676,:$
:7B$>+64-6>;
... Subtribe Ileostylinae Nickrent & Vidal-Russell,
subtr. nov. —
$KG+61$9*#2:'5*8:$K-6D=;$KF@$D676,:1$9*#2:'5*8:$
K-6D=;$F-8=$@76$>+64-6>J$9H$0"+)&%',8:$&Z@@5 ;$E;($K-6D =;J $@E $36 F$
p6:.:7B$:7B$J8#**#)"%&$ K-6D=;$F-8=$E@L,$>+64-6>$67B6C-4$8@$
6:>86,7$*L>8,:.-:;
a,L8-46>$D.:?,-$-7$4:L.-?L>$+:,:>-8-4-$,:B-4-?L>$6+-4@,8-4:O
.-?L>$+,:6B-8-;$a@.-:$@++@>-8:;$<7E.@,6>4678-:$6S$,:46C->$>-C+.-O
4-?L>$A6.$LC?6..->$C@7:B@,LC$8,-:B@,LCA6$4@7>8:7>J$E.@,-?L>$
?,:486->$>@.-8:,-->$>L?8678-?L>;$a.@,6>$?->6SL:.6>$:48-7@C@,+=-J$
:L8$+:,A-$4=@,-+68:.-$^OC6,-$:78=6,->$?:>-E-S->$&9*#2:'5*8:($:L8$
D,:7B6>$D:C@+68:.-$!OC6,-$:78=6,->$B@,>-E-S->$&J8#**#)"%&(;
I.:?,@L>$C->8.68@6>$F-8=$@++@>-86$.6:A6>$:7B$6+-4@,8-4:.$
,@@8>;$K=6$?:>-4$-7E.@,6>46746$L7-8>$:,6$C@7:B>$@,$8,-:B>$:,O
,:7D6B$-7$>-C+.6$,:46C6>$@,$LC?6.>;$<7$J8#**#)"%&$8=6$4678,:.$
E.@F6,$@E$8=6$ 8,-:B$->$>6>>-.6;$3@$?,:48>$:,6$ :>>@4-:86B$F-8=$
8=6$ -7E.@,6>46746J$ ?L8$ 6:4=$ ?->6SL:.J$ :48-7@C@,+=-4$E.@F6,$
->$>L?867B6B$?G$@76$?,:48;$K=6$@A6,:..$E.@,:.$C@,+=@.@DG$@E$
8=6$8F@$D676,:$->$B-EE6,678;$9*#2:'5*8:$=:>$>C:..$&';!$CC(J$
4=@,-+68:.@L>$ E .@F6,>$ F-8=$ E@L,$ +68:.>$ F=6,6:>$ J8#**#)"%&$
=:>$ .:,D6,$ &';!$ 4C(J$D:C@+68:.@L>$E.@F6,>$ F-8=$E-A6$ +68:.>;$
<7$8=6$E@,C6,$D67L>$8=6$:78=6,>$:,6$?:>-E-S6B$:7B$-7$8=6$.:886,$
B@,>-E-S6B;$<7B66BJ$8=6$E.@F6,>$@E$9*#2:'5*8:$,6>6C?.6$P81#"& J$
?L8$8=->$CL>8$=:A6$6A@.A6B$-7$+:,:..6.$D-A67$8=6-,$+=G.@D6768-4$
B->8:746;
... Subtribe Loranthinae Engl. —
$KF@$D67 6,:1 $A#+&)G
)"&$V:,.@F$F-8=$@76$>+64-6>J$AH$2B'8:".2*"&$&T6,,;($V:,.@FJ$@E$
8=6$H=-.-++-76>J$36F$IL-76:J$ 36F$V,-8:-7J$:7B$8=6$Q@.@C@7$
<>.:7B>$:7B$@2)&%',8:$g:4N;$F-8=$4:;$867$>+64-6>$@E$YL,@+6$8@$
>@L8=6,7$`=-7:J$:7B$QLC:8,:;
K=6>6$D.:?,@L>J$:.86,7:86$.6:A6B$C->8.68@6>$6-8=6,$=:A6$&@2G
)&%',8:($@,$.:45$&A#+&))"&($6+-4@,8-4:.$,@@8>;$K=6$-7E.@,6>46746$
=:>$?667$B6>4,-?6B$:>$:$>+-56$E@,$?@8=$D676,:J$?L8$E@,$A#+&)G
)"&J$:BB-8-@7:.$B6>4,-+8-@7>$ 6S->8J$>L4=$:>$ :$ 'OE.@F6,6B$4GC6$
&M:7>6,J$0%"0(J$:$'OE.@F6,6B$LC?6.$&V:,.@F$9$_-67>J$0%X"(J$
@,$:$^OE.@F6,6B$,:46C6$&V:,.@FJ$0%#^(;$K=6>6$B6>4,-+8-A6$B-EO
E6,6746>$=-D=.-D=8$8=6$B-EE-4L.8-6>$:>>@4-:86B$F-8=$-7E.@,6>46746$
86,C-7@.@DG$8=:8$:,6$6>+64-:..G$:4L86$-7$U@,:78=:46:6$&\L-P8J$
0%#0(;$K=6$!O$@,$WOC6,@L>J$4=@,-+68:.@L>$E .@F6,>$:,6$?->6SL:.$
-7$A#+&))"&$?L8$?@8=$?->6SL:.$:7B$L7->6SL:.$E.@F6,>$@44L,$-7$
@2)&%',8:$&+.:78>$ 8=67$B-@64-@L>(;$*78=6,>$:,6$?:>-E-S6B$ -7$
@2)&%',8:$?L8$B@,>-E-S6B$-7$A#+&))"&;
K=6$B@,>-E-S6B$:78=6,>$4:L>6B$T6,,-..$8@$+.:46$8=6$ 8:S@7$
7@F$57@F7$:>$A#+&))"&$-7$-,) 54"*&%',8:;$K=6$D676,-4$7:C6$
-,)54"*&%',8:$Y-4=.6,J$8=:8$ -74.LB6B$:$ =686,@D676@L>$:>>6CO
?.:D6$@E$[.B$:7B$36F$_@,.B$C->8.68@6>J$F:>$,67B6,6B$-..6D-8-O
C:86$:7B$-8>$>+64-6>$:..@4:86B$8@$867$B-EE6,678$D676,:J$-74.LB-7D$
8=6$76F$D67L>$A#+&))"&$&V :,.@F $9$_-67> J$0%X"(;$K=6>6$:L8=@ ,>$
>8:86B1$eA#+&))"&$C:G$? 6$:7$L 7 >+6 4-: . -R6B J$,6.-4$D67L>$7 6:,$8 =6$
>86C$@E$8=->$[.B$_@,.B$.-76;$K=6$@7.G$@8=6,$[.B$_@,.B$C6C?6,>$
@E$8=6$-,)54"*&%',8:$D,@L+$:,6$8=6$E@L,$86C+6,:86$*L>8,:.-:7$
>+64-6>$7@F$,6E6,,6B$ 8@$ J8#**#)"%&;f$K=6>6$4@746+8>$:,6$ 7@8$
-7$4@7E .-48$ F-8=$8=6$ :48L:.$ +=G.@D6768-4$ ,6.:8-@7>=-+>$F=6,6$
9*#2:'5*8:lJ8#**#)"%&J$ A#+&))"&l@2)&%',8:J$:7B$ 8=6$,6C:-7O
-7D$_$j$%$8:S:$:..$ E@,C$>L446>>-A6$4.:B6>;$K=6$7@C674.:8L,:.$
=->8@,G$@E$8=6$D67L>$@2)&%',8:$=:>$?667$C6,4L,-:.;$K=6$7:C6$
674@C+:>>6B$76:,.G$:..$C->8.68@6>$:8$8=6$8-C6$@E$U-77:6L>J$
F:>$>+.-8$-78@$:$+.68=@,:$@E$D676,:$?G$K-6D=6C$&+L?.-4:8-@7>$
?68F667$0#%^$:7B$0%00(J$:7B$,64-,4LC>4,-?6B$?G$M:7>6,$&+L?O
.-4:8-@7>$?68F667$0%'%$:7B$ 0%^0(;$V:,.@F$&0%%!($4@74.LB6B$
8=:8$8=6$7-76$>+64-6>$:>>-D76B$8@$M51,#&)$?G$M:7>6,$F6,6$:..$
4@7>+64-E-4$F-8=$ @H$2C2)&'8:;$_-8=$@H$#8)21&#8:J$8=6$ D67L>$
F:>$8=67$ 8=@LD=8$8@$-74.LB6$PL>8$8F@$ >+64-6>;$ K=6$V*2)&$2.$
A,"%&$8,6:8C678$ &t-L$9$ I-.?6,8J$'//"($.->8>$ >-S$ >+64-6>$ E,@C$
`=-7:$:7B$867$8@8:.$E@,$8=6$D67L>;$T@.64L.:,$6A-B6746$&g6,OT-7D$
ZLJ$+6,>;$4@CC;($>L++@,8>$8=6$C@7@+=G.G$@E$8=6$`=-76>6$:7B$
YL,@+6:7$>+64-6>;
... Subtribe Amyeminae Nickrent & Vidal-Russell,
subtr. nov. —
$KG+6$I05#0&$K-6D=;$3-76$D676,:1$I05#0&$
K-6D=;$ F-8=$ 4:;$ 0//$ >+64-6>$ E ,@C$ T:.6>-:J$ *L>8,:.-:J$ H=-.-+O
+-76>J$:7B$F6>86,7$H:4-E-4b$3&)&',)&%',8:$&\@,8=;($T-N;$F-8=$
8=,66$ >+64-6>$ @E$Q,-$U:75:$ :7B$T:.:G>-:b$3#%',&0"%&$K-6D=;$
F-8=$@76$>+64-6>J$3H$&*5_".2*"&$&a;$TL6..;$6S$V678=;($K-6D=;J$@E$
*L>8,:.-:b$!&+'5*"21,2)&$K-6D=;$F-8=$8=,66$>+64-6>$@E$H:+L:>-:$
:7B$7@,8=6,7$tL667>.:7B$*L>8,:.-:b$!"1*&'"&$K-6D=;$F-8=$8=,66$
>+64-6>$@E$8,@+-4:.$*L>8,:.-:b$!":')"&%',#:$ M:7>6,$F-8=$ @76$
>+64-6>J$!H$02*".*2)&$&\,:L>6($M:7>6,J$@E$36F$IL-76:b$M#*"G
+&%',#:$M:7>6,$F-8=$@76$>+64-6>J$MH$#*&:'"+&$&M6>,;($M:7>6,$@E$
<7B-:b$-&18&%',#:$M :7>6 ,$F-8 =$@7 6$>+ 64-6>J$-H$&*B#)'":""$M:7>6,$
@E$36F$IL-76:b$:7B$724#)"&%',#$M:7>6,$F-8=$E@L,$>+64-6>$E,@C$
6:>86,7$36F$IL-76:$8@$8=6$Q@.@C@7$<>.:7B>;
a,L8-46>$D.:?,-$+L?6>46786>A6$-7$4:L.-?L>$+:,:>-8-4-$,:B-4-O
?L>$6+-4@,8-4:.-?L>$&+,:686,$!"1*&'"&0($+,:6B-8-;$a@.-:$:.86,7:$
@++@>-8:$A6,8-4-..:8:A6;$<7E.@,6>4678-:$6S$,:46C@$LC?6..:$4:+-O
8L.@A6$4@7>8:7>J$E.@,-?L>$-7$8,-:B:$>-C+.-4-:$:DD,6D:8:A6$B->+@O
>-8->J$A6.$:B$E.@,6C$>@.-8:,-LC$,6BL48:b$?,:486->$E.@,:.-?L>$-786,O
BLC$:44,6>4678-?L>$A6.$4L+L.:C$E@,C:78-?L>;$a.@,6>$?->6SL:.6>$
L7->6SL:.6>A6J$!O$A6.$WOC6,-J$4=@,-+68:.-$D:C@+68:.-A6;
I.:?,@L>$@,$ +L?6>4678$C->8.68@6>J$:..$ ?L8$!"1*&'"&$F-8=$
6+-4@,8-4:.$,@@8>$:7B$C@>8$F-8=$@++@>-86$.6:A6>$&:.>@$:.86,7:86$
:7B$A6,8-4-..:86$-7$I05#0&(;$K=6$?:>-4$-7E .@,6>46746$L7-8$:+O
+6:,>$8@$?6$ :$8,-:B$ F=-4=$C:G$ 8=67$?6$:DD,6D:86B$-78@$4@7O
E.@,6>46746>$@E$,:46C6>J$LC?6.>$:7B$4:+-8L.:J$@,$>@C68-C6>$
,6BL46B$8@$>-7D.6$E.@F6,>$@,$:$>-C+.6$LC?6.$@E$BG:B>$-7$>@C6$
I05#0&$&>66$E-D;$!$-7$V:,.@FJ$0%WW(;$<7$!"1*&'"&$:7B$-&18&%G
',#:$8=6$E.@,:.$?,:48>$:,6$67.:,D6BJ$E@.-:46L>J$:7B$4@77:86$8=L>$
4@A6,-7D$8=6$-7E.@,6>46746$BL,-7D$B6A6.@+C678;$K=->$?,:48$ELO
>-@7$->$C@,6$F6:5.G$B6A6.@+6B$-7$>@C6$>+64-6>$@E$724#)"&%',#$
F=6,6$8=6$?,:48>$E@,C$:$4L+L.6$:8$8=6$?:>6$@E$8=6$+6BL74.6;$K=6$
E.@F6,>$:,6$?->6SL:.$&6S46+8$-7$3&)&',)&%',8:(J$D676,:..G$!O$@,$
WOC6,@L>J$4=@,-+68:.@L>$&6;D;J$3&)&',)&%',8:J$!&+'5*"21,2)&J$
!"1*&'"&($@,$D:C@+68:.@L>J$:7B$F-8=$@,$F-8=@L8$,6E.6S6B$.@?6>;$
*7$:,8-4L.:86B$>8G.6$->$>667$-7$!&+'5*"21,2)&J$:$E6:8L,6$,:,6.G$
>667$@L8>-B6$K,-?6$Y.G8,:78=6:6;

!^%
3-45,678$9$:.;$)$Classif ication of SantalalesTA X ON $!%$&'($)$*+,-.$'/0/1$!"#2!!#
Q-S$ D676,:$ ,6+,6>678-7D$ 8=6$ eI05#0&$ 4@C+.6Sf$ F6,6$
>8,@7D.G$>L++@,86B$:>$:$4.:B6$E,@C$C@.64L.:,$:7:.G>6>$&d-B:.O
]L>>6..$9$3-45,678J$'//#:(;$3&)&',)&%',8:J$F6:5.G$>L++@,86B$
:>$>->86,$8@$8=->$4.:B6J$=:>$:$A6,G$B-EE6,678$C@,+=@.@DG1$8=6$
+.:78>$:,6$B-@64-@L>$F-8=$8-7GJ$^OC6,@L>J$L7->6SL:.$E .@F6,>$8=:8$
6C6,D6$E,@C$=@..@F>$-7$8=6$>86C;$d:,-@L>$D676,-4$7:C6>$4@-76B$
?G$K-6D=6C$:7B$M:7>6,$E@,$8:S:$F-8=-7$8=6$I05#0&$4@C+.6S$
&6;D;J$!"+50&%',#:($=:A6$?667$+.:46B$-7$>G7@7@CG$F-8=$I05G
#0&;$K=6$8F@$>+64-6>$@E$I05#0&$>:C+.6B$-7$d-B:.O]L>>6..$9$
3-45,678$&'//#:($F6,6$7@8$C@7@+=G.68-4;$K=6>6$E:48>J$+.L>$8=6$
=-D=$B6D,66$@E$C@,+=@.@D-4:.$A:,-:8-@7$:C@7D$8=6$.:,D6$7LC?6,$
@E$>+64-6>J$=-D=.-D=8$8=6$766B$E@,$:BB-8-@7:.$C@.64L.:,$F@,5$8@$
?6886,$6>8:?.->=$D676,-4$.-C-8>$-7$*CG6C-7:6;
... Subtribe Scurrulinae Nickrent & Vidal-Russell,
subtr. nov. —
$KG+61$7+8))8*&$U;$KF@$D676,:1$7+8))8*&$U;$
F-8=$4:;$!/$>+64-6>$@E$`=-7:J$Q@L8=6:>8$*>-:$:7B$T:.:G>-:b$
:7B$P& _ "**8:$K-6D=;$F-8=$4:;$"!$>+64-6>$@E$8,@+-4:.$*>-:$&<7B-:$
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&\67G:$4@:>8(;
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... Subtribe Dendrophthoinae Nickrent & Vidal-
Russell, subtr. nov. —
$KG+61$!#%C)21,',2#$T:, 8;$a@L ,$D6 76,: 1$
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... Subtribe Emelianthinae Nickrent & Vidal-Russell,
subtr. nov. —
$KG+6$O0#*"&%',#$M:7>6,;$Q6A67$D676,:1$O0#*"G
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... Subtribe Tapinanthinae Nickrent & Vidal-Russell,
subtr. nov. —
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. Family Opiliaceae Valeton (). —
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. Family Comandraceae Nickrent & Der, fam. nov. —
$
KG+61$A20&%C)&$3L88;$KF@$D676,:1$A20&%C)&$3L88;$F-8=$@76$
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. Family Thesiaceae Vest (). —
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. Family Cervantesiaceae Nickrent & Der, fam. nov.
—
$
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:$F=-86J$E.6>=G$-776,$6S@4:,+$4@A6,-7D$@7$8=6$>8@7G$C6>@4:,+;

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TA XO N$!%$&'($)$*+,-.$'/0/1$!"#2!!#3-45,678$9$:.;$)$Classif ication of Santalales
K=6$4.:B6$8=:8$-74.LB6>$8=6$D676,:$@E$`6,A:786>-:46:6$->$
>8,@7D.G$>L++@,86B$?G$C@.64L.:,$B:8:$:7B$-786,D676,-4$,6.:8-@7O
>=-+>$:,6$EL..G$,6>@.A6B$&M6,$9$3-45,678J$'//#b$]@D6,>$9$:.;J$
'//#(;$K=6$E:C-.G$ ->$ 4@C+@>6B$@E$8F@$4.:B6>J$ 8=6$ E-,>8$ F-8=$
I+&%',2:5)":J$A#)<&%'#:"&$:7B$e2C"%&$&8=6$Q@L8=$*C6,-4:7$@,$
A#)<&%'#:"&$ 4.:B6($:7B$:$ >64@7B$4@C+@>6B$@E$8=6$ ,6C:-7-7D$
E@L,$ D676,:$ &-5)8*&)"&$4.:B6(;$V-@D6@D,:+=-4:..GJ$-5)8*&)"&$
>=@F>$8=6$6:>86,7$3@,8=$*C6,-4:7J$6:>86,7$*>-:7$B->PL748-@7$
:.>@$>667$-7$38+\*#5&$&K=6>-:46:6(;$K=6$.:,D6$E,L-8>$@E$I+&%G
',2:5)":$&%%2%&48:'&'&$`;$i..@:$9$H;$gu,D$:,6$6:867$?G$.@4:.$
+6@+.6;$H:,:>-8->C$@E$>L,,@L7B-7D$A6D68:8-@7$?G$IH$&:"1&12'#$
T;$366$:7B$N\28B&\&$=:>$?667$B@4LC6786B$:7B$->$>LEE-4-678.G$
>6A6,6$8@$.-C-8$D,@F-7D$4,@+>$:,@L7B$8=6$+.:78>;
. Family Nanodeaceae Nickrent & Der, fam. nov. —
$
KG + 61 $ S& %2C#&$ V:75 >$ 6S$ `;a;$I :6,87;$ KF@$ D676,:1$ J"C&$
*;$`L77;$6S$Y7B.;$F-8=$@76$>+64-6>J$JH$ :&*"+".2*"&$$$*;$`L77;J$
B->PL748$E,@C$36F$p6:.:7B$8@$8=6$gL:7$a6,7:7B6R$<>.:7B>b$:7B$
S&%2C#&$V:75>$6S$`;a;$I:6,87;$F-8=$@76$>+64-6>J$SH$08:+2:&$
`;a;$I:6,87;J$@E$86C+6,:86$ Q@L8=$*C6,-4:$&H:8:D@7-:J$K-6,,:$
B6.$aL6D@J$:7B$8=6$a:.5.:7B$<>.:7B>(;
*,?@,6>$&J"C&($A6.$ >LEE,L8-46>$ &S&%2C#&($ -7$ ,:B-4-?L>$
+:,:>-8-4-;$ a@.-:$:.86,7:J$867L-:$ 4:,7@>:A6;$a.@,6>$?->6SL:.6>$
^OC6,-$ 4:.G4L.@$>L?8678-J$B->4-$.@?L.->$D.:7BL.:,-?L>$-786,$86O
+:.:$:.86,7:78-?L>J$:78=6,:,LC$4@77648-A->$:B$86+:.:$8,-4=@C:O
8-?L>$:EE-S->J$@A:,-@$>6C-O-7E6,@$&J"C&($A6.$-7E6,@$&S&%2C#&(;$
a,L48L>$6S$B,L+:$+6,-:78=-@$+6,>->86786$4@,@7:8:$4@7>8:7>;
J"C&$->$:$=6C-+:,:>-8-4$8,66$F=6,6:>$S&%2C#&$->$:$8L,EO$@,$
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:7B$>@C6F=:8$E.6>=G;$K=6$E .@F6,>$:,6$?->6SL:.J$868,:C6,@L>J$
F-8=$:$4:.G4L.L>;$Z:-,>$@++@>-86$8=6$>8:C67>$:>$F6..$:>$:$.@?6B$
D.:7BL.:,$B->5$:,6$ +,6>678;$K=6$@76O.@4L.:,$@A:,G$->$-7E6,-@,$
-7$S&%2C#&$:7B$=:.E$-7E6,-@,$-7$J"C&;$K=6$+.:4678:.$4@.LC7$->$
>8,:-D=8$:7BJ$:8$.6:>8$-7$J"C&$6S867B>$?6G@7B$8=6$@AL.6>$:+-4:..G$
:>$:$4@7-4:.$+,@P648-@7$ 8@$ 8=6$ ?:>6$@E$ 8=6$ >8G.6;$KF@$@,$8=,66$
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4,@F76B$?G$8=6$+6,>->8678$+6,-:78=$+:,8>;
J"C&$+,6>678>$D676,:.-R6B$A6D68:8-A6$:7B$E.@,:.$4=:,:486,>J$
8=L>$-8$->$7@8$>L,+,->-7D$8=:8$-8$F:>$4.:>>-E-6B$F-8=$C6C?6,>$@E$
Q:78:.6:6$>L4=$:>$V8:&%8:$:7B$O8+&)5&J$?@8=$7@F$-74.LB6B$-7$
7&%'&*80$&H-.D6,J$0%"!(;$K=->$ ,6.:8-@7>=-+$F:>$>L++@,86B$ ?G$
6C?,G@.@D-4:.$E6:8L,6>J$>L4=$:>$8=6$-2*542%80$6C?,G@$>:4$8=:8$
6S867B>$?6G@7B$8=6$@AL.6$-7$8=6$^O7L4.6:86$>8:D6;$VL8$A:,-:8-@7>$
@7$8=->$e:DD,6>>-A6f$6C?,G@$>:4$?6=:A-@,$:.>@$@44L,>$-7$@8=6,$
Q:78:.:46:6$>;.;$:>$F6..$:>$-7$[+-.-:46:6J$T->@B67B,:46:6$:7B$
U@,:78=:46:6J$8=L>$8=->$E6:8L,6$4:77@8$?6$L>6B$8@$.-75$J"C&$8@$
:7G$+:,8-4L.:,$>:78:.:.6:7$4.:B6;$K=6$D6@D,:+=-4:.$B->8,-?L8-@7>$
@E$8=6>6$8F@$D676,:$>LDD6>8>$8=6G$C:G$?6$I@7BF:7:7$,6.-48>;
. Family Santalaceae R. Br. (), emend. —
$Y.6A67$
D676,:1$I%'"C&1,%#$H@6++;$9$Y7B.;$F-8=$>6A67$>+64-6>$@E$Q@L8=$
*C6,-4:J$8=6$`:,-??6:7J$:7B$T6S-4@b$A2*122%$H; g ;$ V 6 , D - L >$ q - 7 O
4.LB-7D$V8:&%8:$+201)#::8:$TL,,r$F-8=$8F@$>+64-6>$@E$Q@L8=$
*E,-4:b$O8B)&+,"2%$Z@@5;$E;$F-8=$8F@$>+64-6>$E,@C$8=6$`:,-?O
?6:7$8@$Q@L8=$*C6,-4:b$O_2+&)12:$U:?-..;$F-8=$'W$>+64-6>$@E$
Q@L8=6:>8$*>-:$:7B$T:.:G>-:$8@$Z:F:--b$@#1"C2+#)&:$Z@@5;$E;$
F-8=$8F@$>+64-6>$@E$`=-.6b$J52:+,"*2:$]L-R$9$H:A;$F-8=$ @76$
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M`;$F-8=$ @76$>+64-6>b$NH$ &+#)B&$ &];$V,;($*;$M`;J$@E$>@L8=O
6:>86,7$*L>8,:.-:b$N:5)":$U;$F-8=$8F@$>+64-6>$ @E$YL,@+6J$8=6$
T6B-86,,:76:7J$*E,-4:$8@$`=-7:b$K,2"&+&)12:$*;$M`;$F-8=$@76$
>+64-6>J$KH$+&1#%:":$&Z:,A;($*;$M`;J$67B6C-4$8@$Q@L8=$*E,-4:b$
:7B$7&%'&*80$U;$F-8=$0W$>+64-6>$E,@C$<7B@C:.:G:$8@$*L>8,:.-:$
:7B$Z:F:--;$<74.LB6>$Y,6C@.6+-B:46:6$K-6D=;$6S$\L-P8;
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&C->8.68@6>(;$U6:E$E@,C$->$A:,-:?.6$F-8=$:.86,7:86$:7B$@++@>-86$
:,,:7D6C678>J$+6,>->8678$@,$4:BL4@L>J$6S+:7B6B$.:C-7:$@,$,6O
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-74.LB-7D$?->6SL:.$E.@F6,>J$B-@64-@L>J$C@7@64-@L>J$:7B,@C@7O
@64-@L>J$:7B$:7B,@B-@64-@L>;$ <7E.@,6>46746>$:,6$:S-..:,G$:7B$
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>+-56>J$4GC6>$&>-C+.6$:7B$4@C+@L7B$B-4=:>-:(J$+:7-4.6>J$E:>O
4-4.6>J$:7B$LC?6.>;$*$4:.GS$->$:?>678$@,J$-7$J52:+,"*2:J$+,6>678$
:>$:$4:.G4L.L>;$H68:.>$"2WJ$E,66$@,$EL>6B$-78@$:$4@,@..:$8L?6$8=:8$
->$>=@,8$:7B$4:C+:7L.:86$@,$L,46@.:86;$H68:.$=:-,>$@++@>-86$8=6$
>8:C67>$C:G$?6$+,6>678$@,$:?>678$&O_2+&)12:J$N01,&+20#)"&J$
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8@$-7E6,-@,$&6;D;J$A2*122%(;$<7$>@C6$D676,:J$>L4=$:>$A2*122%J$
K,2"&+&)12:$:7B$N:5)":$8=6$.@4L.6$ ->$ .@?6B$ :8$ 8=6$?:>6$8=L>$
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>@C68-C6>$F-8=$8=6$>L?867B-7D$+6B-46.$>F@..67$:7B$E.6>=G$&6;D;J$
O_2+&)12:(;$<7$8=6$:6,-:.$+:,:>-86>$8=6$E.6>=G$C6>@4:,+J$E@,C>$
E-?,@L>$>66B$:88:4=C678$>8,L48L,6>;
Q:78:.:46:6$>;>8,;$F6,6$=-D=.G$>L++@,86B$:>$C@7@+=G.68-4$
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4.:B6$4@78:-7-7D$8=,66$D676,:$@E$36F$_@,.B$C->8.68@6>$F=@>6$
C6C?6,>$=:A6$?667$:>>-D76B$8@$:$A:,-68G$@E$>:78:.:.6:7$E:C-O
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0%##( J$ Q: 78 :.: 46 :6$ & _-6 7> $ 9$ V: ,.@ FJ$ 0%X0(J$ : 7B $ d-> 4:4 6: 6$
&V=:7B:,-$9$d@=,:J$0%#"(;$K,-?6$*78=@?@.6:6J$E@,C6,.G$4@7O
8:-7-7D$I%',2B2*8:J$N01,&+20#)"&$:7B$O_2+&)12:J$->$ 7@F$
B6EL748$D-A67$8=:8$I%',2B2*8:$=:>$ ?667$ 8,:7>E6,,6B$8@$[+-.-O
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8=6$,6C:-7-7D$D676,:$-7$8=6$E:C-.G;
. Family Amphorogynaceae (Stauffer ex Stearn) Nick-
rent & Der, stat. nov. —
$V:>6B$@7$8,-?6$*C+=@,@DG76:6$
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9$Q8:LEE;$F-8=$@76$>+64-6>J$!H$+2)&**"%&$Zv,.;J$67B6C-4$8@$36F$
`:.6B@7-:b$!#%C)205/&$M:7>6,$q-74.LB-7D$A*&C205/&$M: 7>6,r $
F-8=$'0$>+64-6>$@E$>@L8=6:>86,7$*>-:J$T:.:G>-:J$<7B@76>-:J$:7B$
36F$IL-76:b$!#%C)2')21,#$T-N;$q-7 4.LB-7D$M#%:*2c"&$V.LC6r$
F-8=$E@L,$>+64-6>$E,@C$ 8=6$ Z-C:.:G:>$8@$8=6$H=-.-++-76>$ :7B$
T:.:G>-:J$<7B@76>-:b$!8.)#%25&$`=:8-7$F-8=$6.6A67$>+64-6>$@E$
<7B@76>-:$:7B$T:.:G>-:b$@#1'20#)"&$];$V,;$F-8=$0X$>+64-6>$@E$

!!"
3-45,678$9$:.;$)$Classif ication of SantalalesTA X ON $!%$&'($)$*+,-.$'/0/1$!"#2!!#
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:7B$>86CO+:,:>-8-4($>=,L?>J$>86C$B67B,@+:,:>-86>$&.-:7:>(J$:7B$
>86CO+:,:>-8-4$>=, L?>$&C->8.68@6>(;$-,&+#**&)"&$->$:$=G+6,+:,:O
>-86$@7$U@,:78=:46:6$:7B$@8=6,$*C+=@,@DG7:46:6;$Q86C>$F-8=$
>GC+@B-:.$?,:74=-7DJ$-7$>@C6$:6,-:.$+:,:>-86>$B-EE6,678-:86B$-78@$
A6D68:8-A6$:7B$=:L>8@,-:.$>=@@8>;$U6:A6>$:.86,7:86$@,$>L?A6,8-O
4-..:86J$6-8=6,$B-C@,+=-4$@,$7@8$B-C@,+=-4J$+6,>->8678J$4:BL4@L>$
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46746>$A:,-@L>$-74.LB-7D$>@.-8:,G$E.@F6,>J$C@7@4=:>-:J$>+-56>J$
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>8:C67>$:,6$+,6>678$-7$C@>8$D676,:$&!#%C)205/&$:7B$-,&+#*G
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6NL:.$8@$@,$>.-D=8.G$>=@,86,$@,$.@7D6,$8=:7$8=6$:78=6,;$K=6$:78=6,>$
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8=6$:78=6,$->$>6>>-.6;$<7>86:B$@E$8=6$.@4L.6>$@E$8=6$868,:>+@,:7D-:86$
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>@,$->$+,6>678;$K=6$C:8L,6$E,L-8$&B,L+6($->$4,@F76B$?G$+6,>->8678$
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TA XO N$!%$&'($)$*+,-.$'/0/1$!"#2!!#3-45,678$9$:.;$)$Classif ication of Santalales
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PERSPECTIVES ON THE NEW SANTALALES
CLASSIFICATION
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8=6$E:C-.-6>$4@78:-7$@7.G$8F@$@,$8=,66$D676,:;$<7$C@>8$@E$8=6>6$
E:C-.-6>J$ >6A6,:.$@E$8=6$4@C+@7678$D676,:$:,6$:.>@$C@7@>+6O
4-E-4J$ 8=L>$>LDD6>8-7D$8=6>6$ 4.:B6>$,6+,6>678$ ->@.:86B$:7Bl@,$
,6.-48L:.$4.:B@D67-4$6A678>;
_6$?6.-6A6$8=:8$ 8=6$ 4.:B6>$ @?8:-76B$E,@C$CL.8-D676$C@O
.64L.:,$:7:.G>6>$,6+,6>678$8=6$?6>8$4L,,678$=G+@8=6>6>$E@,$8=6$
6A@.L8-@7:,G$.-76:D6>$F-8=-7$Q:78:.:.6>;$a,@C$8=6>6$4.:B6>$F6$
=:A6$+,@+@>6B$:$,6A->6B$4.:>>-E-4:8-@7$@E$8=6$@,B6,J$G68$8=6$,6O
>L.8-7D$E:C-.-6>$:,6J$-7$>@C6$4:>6>J$C@,6$B-EE -4L.8$8@$-B678-EG$
:7B$ 4-,4LC>4,-?6$ L>-7D$C@,+=@.@D-4:.$E6:8L,6>;$ 3LC6,@L>$
6S:C+.6>$4:7$ ?6$ D-A67$ F=6,6$ C@.64L.:,$+=G.@D6768-4$C68=O
@B>$=:A6$-B678-E-6B$:$>8,@7D.G$>L++@,86B$4.:B6$8=:8$.:45>$C@,O
+=@.@D-4:.$>G7:+@C@,+=-6>$&`=:>6$ 9$:.;J$'///b$QC-8=$9$:.;J$
'//W(J$?L8$,64@D7-8-@7$@E$8=6$D,@L+$ ->$ PL>8-E-6B$ ?G$:>>LC-7D$
>L4=$>G7:+@C@,+=-6>$&?6$8=6G$C-4,@C@,+=@.@D-4:.J$4=6C-4:.$
@,$D67@C-4($6S->8$ :7B$ ,6C:-7$ 8@$ ?6$ B->4@A6,6B;$_6$+,6>678$
?6.@F$:$56G$8=:8$L>6>$:A:-.:?.6$C@,+=@.@D-4:.$-7E@,C:8-@7$8@$
-B678-EG$8=6$0#$E:C-.-6>$@E$Q:78:.:.6>;$Y>+64-:..G$E@,$[.:4:46:6$
>;.;$:7B$Q:78:.:46:6$>;.;J$:7:8@C-4:.$:7B$6C?,G@.@D-4:.$E6:8L,6>$
F6,6$>@C68-C6>$6C+.@G6B$?64:L>6$C@>8$C:4,@C@,+=@.@D-4:.$
E6:8L,6>$:,6$ +@.GC@,+=-4$ :C@7D$ 4.:B6>;$`=:,:486,>$ >L4=$ :>$
+:,:>-8->C$C@B6$&>86C$A>;$,@@8(J$+.:78$>6SL:.$4@7B-8-@7J$@A:,G$
+@>-8-@7J$+68:.$=:-,$8LE8>$@++@>-86$>8:C67>J$>66B$A->4-7$8->>L6J$
684;$=:A6$>8:86>$8=:8$:++6:,$CL.8-+.6$8-C6>$-7$B-EE6,678$4.:B6>;$
*>$B->4L>>6B$-7$M6,$9$3-45,678$&'//#(J$-8$>66C>$8=->$A:,-:8-@7$

!!!
3-45,678$9$:.;$)$Classif ication of SantalalesTA X ON $!%$&'($)$*+,-.$'/0/1$!"#2!!#
Agarwal, S. 0%W";$T@, +=@.@D-4:.$: 7B$6C? ,G@.@D-4: .$>8L B- 6>$-7$8=6 $E:CO
-.G$[.:4:46:6;$<;$N*&_$U;$-,5'202)1,2*24 5$0"1$0#!20%W;
APG (Angiosperm Phylogeny Group). 0%%#;$*7$@,B-7:.$ 4.:>>-E-4:O
8-@7$E@,$8=6$E:C-.-6>$@E$E.@F6,-7D$+.:78>;$I%%H$J"::28)"$32'H$L&)CH$
#!1$!"02!!";
APG (Angiosperm Phylogeny Group). '//";$*7$L+B:86$@E$8=6$*7D-@O
>+6,C$H=G.@D67G$I,@L+$4.:>>-E -4:8-@7$E@,$8=6$@,B6,>$:7B$E:C-.-6>$
@E$E.@F6,-7D$+.:78>1$*HI$<<;$32'H$eH$@"%%H$72+H$0^01$"%%2^"W;
APG (Angiosperm Phylogeny Group). '//%;$*7$L+B:86$@E$8=6$*7D-@O
>+6,C$H=G.@D67G$I,@L+$4.:>>-E -4:8-@7$E@,$8=6$@,B6,>$:7B$E:C-.-6>$
@E$E.@F6,-7D$+.:78>1$*HI$<<<;$32'H$eH$@"%%H$72+H$0W01$0/!20'0;
Ashworth, V.E.T.M. '///:;$ H=G.@D6768-4$,6.:8-@7>=-+>$ -7$ H=@,:B67O
B,6:6$&d->4:46:6($-7E6, ,6B$E,@C$8= ,66$,6D-@7>$@E$8=6$7L4.6:,$,-?@O
>@C:.$4->8,@71$<;$T:P@,$.-76:D6>$:7B$+:,:+=G.G$@E$-,2)&C#%C)2%;$
75:'H$32'H$'!1$"^%2"X/;
Ashworth, V.E.T.M. '///?;$H=G.@D6768-4$,6.:8-@7>=-+>$ -7$ H=@,:B67O
->$E,:48:.$-7$7:8L,6J$-7$@8=6,$F@,B>J$>-C-.:,$8=6C6>$:++6:,$,6O
+6:86B$:8$B-EE6,678$=-6,:,4=-4:.$.6A6.>;$<7$D676,:.J$L7:C?-DL@L>$
>G7:+@C@,+=-6>$ B6.-C-8-7D$E:C-.-6>$:7B$.:,D6,$ D,@L+-7D$:,6$
,:,6J$8=L>$ 4=:,:486,-R:8-@7$ L>L:..G$,6NL-,6>$L7-NL6$ 4@C?-7:O
8-@7>$@E$4=:,:486,>;
KEY TO SANTALALES FAMILIES
$ 0;$ H:,:>-8-4$@,$7@8b$E .@F6,>$B-4=.:CGB@L>$&6S46+8$>@C6$N+G
'2\%#0&$@E$[48@576C:46:6(b$>8:C67>$6NL:.$8@$@,$D,6:86,$
-7$7LC?6,$8=:7$+68:.>b$@AL.6>$L7-86DC-4$@,$?-86DC-4b$>66B>$
F-8=$:$8=-7$86>8:;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$ '
$ 0;$ H:,:>-8-4b$E.@F6,>$C@7@4=.:CGB@L>$&>6+:.>$:?>678($@,$B-O
4=.:CGB@L>$&@E867$F-8=$,6BL46B$>6+:.>(b$>8:C67$:7B$+68:.$
7LC?6,>$6NL:.b$@AL.6>$:86DC-4b$>66B>$F-8=@L8$:$86>8:;$;$ #
$ ';$ Q86..:86$+L?6>46746$+,6>678b$E.@F6,>$L7->6SL:.$&+.:78>$B-O
@64-@L>(b$@A:,G$-7E6,-@,;$;$;$;$;$;$;$;$;$;$;$;$;$;Octoknemaceae
$ ';$ Q86..:86$+L?6>46746$:?>678b$E.@F6,>$?->6SL:.$&6S46+8$M&)G
0&%C"&$:7B$M2%C8)2C#%C)2%$@E$*+8 : 7B,:4 6:6(b$@A:, G$>LO
+6,-@,$&6S46+8$>@C6$7')20B2:"&$@E$Q8,@C?@>-:46:6( ;$;$;$"
$ ";$ 3@8$+:,:>-8-4$&=:L>8@,-:$:?>678($$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$^
$ ";$ ]@@8O+:,:>-8-4$&=:L>8@,-:$+,6>678(;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$W
$ ^;$ U6:E$ B67B,-8-4$ =:-,>$+,6>678b$ 6+-B6,C:.$ 46..>$.-D7-E-6Bb$
.6:A6>$F-8=$>4=-R@D67@L>$4:A-8-6>b$>8:C67$ F=@,.>$8F@$@,$
8=,66 ;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$ Coulaceae
$ ^;$ U6:E$B67B,-8-4$=:-,>$:?>678b$6+-B6,C:.$46..>$7@8$.-D7-E-6Bb$
.6:A6>$F-8=@L8$>4=-R@D67@L>$4:A-8-6>b$>8:C67$F=@,.>$@76$
@,$8F@$$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$!
$!;$Y+-B6,C:.$46..$B,L>6>$+,6>678$&6S46+8$7+2)2C2+&)18:(b$
>6+:.>$4@C+.686.G$4@77:86$$;$;$;$;$;$;$;$;$;$;$ Strombosiaceae
$ !;$ Y+-B6,C:.$46..$B,L>6>$:?>678b$>6+:.>$+:,8-:..G$4@77:86;$;$;
;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$; Erythropalaceae
$ W;$ <7E .@,6>46746$:$>-C+.6$LC?6.b$4:.GS$7@8$:44,6>4678$$$;$;$;$;
;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$Ximeniaceae
$ W;$ <7E .@,6>46746>$A:,-@L>$?L8$7@8$:$>-C+.6$LC?6.b$4:.GS$:4O
4,6>4678$@,$7@8 ;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$ X
$ X;$ Q8:C67>$-7$@76$F=@,.b$:78=6,$B6=->46746$?G$+@,6>$@,$E.:+>b$
@A:,G$0O$@,$'O.@4L.:, ;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$Aptandraceae
$ X;$ Q8:C67>$&-74.LB-7D$>8:C-7@B6>($-7$8F@$F=@,.>b$:78=6,$B6O
=->46746$?G$>.-8>b$@A:,G$0O.@4L.:,$$$;$;$;$;$;$;$;$;$;$ Olacaceae
$ #;$ Q86CO+:,:>-8-4b$E,L-8$:7$:4=676$?6:,-7D$:44,6>4678$E6:8=6,G$
>8:C-7@B6>$:.86,7:86$F-8=$+6,-:78=$.@?6> ;$;$;$;$;$;$;$;$;$;$;$;$;$
;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$Misodendraceae
$#;$Q86CO$@,$,@@8O+:,:>-8-4b$E,L-8$:$B,L+6J$0O>66B6B$?6,,GJ$7L8.68$
&7@8$:$+.LC@>6$:4=676($$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$%
$%;$M->8G.@L>b$E.@,:.$?,:48>$:7B$?,:486@.6>$EL>6B$-78@$:$4L+$
-7$W8"%+,&0&*"80$:7B$7+,2#1."&b$+68:.>$ 4@77:86$ -78@$:$
8L?6b$+68:.$ =:-,>$ @++@>-86$ 8=6$ >8:C67>$ +,6>678$-7$I)b2%&$
:7B$7+,2#1."& ;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$Schoepfiaceae
$ %;$ 3@8$ B->8G.@L>b$ E .@,:.$ ?, :48>$ :7B$ ?,: 486@.6>$ &-E$ +,6>678($
>6+:,:86b$ +68:.>$E,66$ @,$ 4@77:86b$+68:.$ =:-,>$@++@>-86$8=6$
>8:C67>$+, 6>678$@,$:?>678 ;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$ 0/
0/;$ *786,-@,$:7B$+@> 86,-@,$:78 =6,$.@4 L.6>$@E$L 76NL :.$>-R6J$6:4=$
8=64LC$B6=->4-7D$8,:7>A6,>6.G$-7B6+67B678.Gb$+68:.>$F-8=$
L74-7:86$8-+> ;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$Amphorogynaceae
0/;$ *786,-@,$:7B$+@>86,-@,$:78=6,$.@4L.6>$@E$6NL:.$>-R6J$6:4=$
8=64LC$B6=->4-7D$?G$:$4@CC@7$.@7D-8LB-7:.$>.-8$@,$&-7$d":G
+80($:78=6,>$EL>6B$F-8=$+@,@>6$B6=->46746b$+68:.>$F-8=@L8$
L74-7:86$8 -+>$$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$ 00
00;$ U6:E$4G>8@.-8=>$+,6>678$&:?>678$-7$I%',2B2*8:(b$@A:,G$>LO
+6,-@,;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$Opiliaceae
00;$ U6:E$4G>8 @.-8=>$:?>678b$@A:,G$=:.E$@,$E L..G$-7E6, -@,$$;$;$;$ 0'
0';$ `:.G S$+,6>678$:>$E@.-:46L>$>6+:.>$&E6C:.6$E.@F6,>$@E$38+\G
*#5&($@,$:$4:.G4L.L>$&4:.GS$,6C7:78( ;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$ 0"
0';$ `:.GS$@,$4:.G4L.L>$:?>678 ;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$ 0!
0";$ ]@@8$:7B$>86C$+:,:>-86>b$+68:.$=:-,>$@++@>-86$>8:C67>$:?O
>678b$>66B$F-8=$A->4-7$8->>L6 ;$;$;$;$;$;$;$;$;$;$;$ Loranthaceae
0";$ ]@@8$+:,: >-86>b$+68:.$=:-,>$@++@>-86$>8:C67>$+,6>678$&:?>678$
-7$38+\*#5&(b$>66B$F-8=@L8$A->4-7$8-> >L6;$;$;$;$;$;$;$;$;$;$;$;$ 0^
0^; $ a ,L -8 $ C6 >@ 4: ,+ $ E .6 >= GJ$ 6 7B @4: , +$ F :. .$ ? @7G $ @, $ 4: , 8- .: D- O
7@L>;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$; Nanodeaceae
0^; $ T6>@4: , +$ ?@7GJ$ 67B@4:, + $ + : , 6 74 = G C :8 @ L >$ @ , $ 4 , L> 8 : O
46@L> ;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$; Thesiaceae
0!;$ YC?,G@$>L>+67>@,$+,6>678$$;$;$;$;$;$;$;$;$;$;$;$;$;$; Santalaceae
0!;$ YC?, G@$>L>+67>@,$:?>678 ;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$ 0W
0W; $ Q86CO + : , : >-8 - 4b$+= G..@ 8 : SG$B64L> > : 86b $> 6 6 B$F -8=$A-> 4 - 7$8 ->O
>L6 ;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$; Viscaceae
0W;$ ]@@8O+:,:>-8-4b$ +=G..@8:SG$ :.86,7:86b$>66B$F-8=@L8$A->4-7$
8->>L6 ;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$ 0X
0X;$ Q86C>$ :7B$.6:A6>$+L?6>4678b$ 8,66>$@,$>=,L?>$$$;$;$;$;$;$;$;$;$;$
;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$Cervantesiaceae
0X;$ Q86C>$ :7B$.6:A6>$D.:?,@L>b$=6,?:46@L>$+6,677-:.>$$;$;$;$;$;$
;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$;$ Comandraceae
ACKNOWLEDGEMENTS
K=6$C@.64L.:,$+=G.@D6768-4$>8LB-6>J$F=-4=$E@,C$8=6$E@L7B:8-@7$
@E$8=->$F@,5J$,6+,6>678$8=6$4L.C-7:8-@7$@E$6EE@,8>$?G$C:7G$-7B-A-BL:.>$
-7$ :BB -8-@7$8@$8=6$ 4@:L8=@,>;$ K=6>6$ -74.LB6$ 4@..6:DL6>$ :7B$>8LB678>$
F=@$ :>>->86B$-7$ .:?@,:8@, G$ F@,5J$ >4-678->8>$ F=@$ 4@..6486B$ :7B$ >678$
+:,: >-8-4$+.:78$>:C+.6>J$:7B$8=@>6$F=@$+,@A-B6B$:>>->8:746$F-8=$E-6.B$
F@,5$:7B$B:8:$:7:.G>6>;$ _6$:,6$+:,8 -4L.:,.G$ D,:86E L.$ 8@$]@G$I6,6:L$
&T->>@L,-$V@8:7-4:.$I: ,B67($F=@$+,@A-B6B$U:8-7$8,:7>.:8-@7>$@E$8=6$
76F$8:S@7$B6>4,-+8-@7>;
LITERATURE CITED

!!W
TA XO N$!%$&'($)$*+,-.$'/0/1$!"#2!!#3-45,678$9$:.;$)$Classif ication of Santalales
B,6:6$&d->4:46:6($-7E6, ,6B$E,@C$8= ,66$,6D-@7>$@E$8=6$7L4.6:,$,-?@O
>@C:.$4->8,@71$<<;$K=6$3@,8=$*C6,-4:7$>+64-6>$@E$-,2)&C#%C)2%;$
I*":2$0%1$^02!";
Baas, P. 0%#';$U6:E$:7:8@CG$:7B$ 4.:>>-E -4:8-@7$@E$8=6$[.:4:46:6J$N+G
'2\%#0&$:7B$O)5',)21&*80;$I**#)'2%"&$"1$0!!2'0/;
Backlund, A. & Bremer, K. 0%%#;$K@$?6$ @,$ 7@8$ 8@$ ?61$ H,-74-+.6>$ @E$
4.:>>-E-4:8-@7$:7B$C@7@8G+-4$+.:78$E:C-.-6>;$P&_2 % $^X1$"%02^//;
Baillon, H.M. 0#W';$M6LS-{C6$CcC@-,6$>L,$.6>$U@,:78=:4c6>;$IC&%G
:2%"&$"1$!/20'#;
Barkman, T.J., McNeal, J.R., Lim, S.-H., Coat, G., Croom, H.B.,
You ng , N .D . & de Pa mp hi li s, C.W . '//X;$T-8@4=@7B,-:.$M3*$>LDO
D6>8>$:8$.6:>8$00$@,-D-7>$@E$+:,:>-8->C$-7$:7D-@>+6,C>$:7B$,6A6:.>$
D67@C-4$4=-C6,->C$-7$+:,:>-8-4$+.:78>;$3JA$O<2*H$3"2*H$X1$'^#;
Barlow, B.A. 0%WW;$*$,6A->-@7$ @E$8=6$ U@,:78=:46:6$@E$*L>8,:.-:$:7B$
36F$p6:.:7B;$I8:')&*H$eH$32'H$0^1$^'02^%%;
Barlow, B.A. 0% #";$ V-@ D6@ D, :+= G$ @E$ U @,: 78 =:4 6: 6$ :7 B$ d-> 4: 46: 6;$
H+;$0%2^!$-71$`:.B6,J$M;T;$9$V6,7=:,B8J$H;$&6B>;(J$P,#$B"2*245$2.$
0":'*#'2#:;$36F$n@,51$*4:B6C-4$H,6>>;
Barlow, B.A. 0%#^;$U@,:78=:46:6;$H+;$W#20"0$-71$I6@,D6J$*;Q;$&6B;(J$
V*2)&$2.$I8:')&*"&;$`:7?6,,:1$*L>8,:.-:7$I@A6,7C678$HL?.->=-7D$
Q6,A-46;
Barlow, B.A. 0%%0;$`@7>+648 L>$@E$8=6$D676,:$7+8))8*&$U;$:7B$P&_"* *8:$
K-6D=6C$&U@,:78=:46:6($-7$8=6$T:.6>-:7$,6D-@7;$3*80#&$"W1$W"2# !;
Barlow, B.A. 0%%!;$36F$ :7B$ 7@86F@,8=G$T:.6>-:7$>+64-6>$ @E$U@,:7O
8=:46:6;$3*80#&$^/1$0!2"0;
Barlow, B.A. & Wiens, D. 0% X";$ K =6$ 4. :>> -E -4: 8-@ 7$ @E$ 8 =6$ D6 76 ,-4 $
>6D,6D:86>$@E$-,)54"*&%',8:$&j$S2'&%',#)&($@E$8=6$U@,:78=:46:6;$
3)"''2%"&$'!1$'W2"%;
Bentham, G. & Hooker, J.D. 0##";$L#%#)&$1*&%'&)80;$U@7B@71$]66A6;
Beyer, C., Forstreuter, W. & Weber, H.C. 0%#%;$*7:8@C-4:.$>8LB-6>$
@E$=:L>8@,-L C$@78@D67G$ :7B$ ,6C:,5:?.6$ C@B6$ @E$ +6768,:8-@7$@E$
8=6$=:L>8@,-LC$-7$S85':"&$ .*2)"B8%C&$&U:?-..;($];$V,;$32'H$ I+'&$
0/'1$''%2'"!;
Bhandari, N.N. & Vohra, S.C.A. 0%#";$YC?,G@.@DG$:7B$:EE-7-8-6>$@E$
d->4:46:6;$H+;$W%2#W$-71$`:.B6,J$ T;$9$V6,7=:,B8J$ H;$&6B>;(J$P,#$
B"2*245$2.$0":'*#'2#:;$36F$n@,51$*4:B6C-4$H,6>>;
Bhatnagar, S.P. & Johri, B.M. 0%#";$YC?,G@.@DG$@E$U@,:78=:46:6;$
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V. , C h as e , M . , Po we l l, M . , A l i ce , L ., E va n s, R . , S au q ue t , H . ,
Neinhuis, C., Slot ta, T., Rohwer, J., Campbell, C. & Chatrou,
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3-45,678$9$:.;$)$Classif ication of SantalalesTA X ON $!%$&'($)$*+,-.$'/0/1$!"#2!!#
Lee, S.-K. 0%#/;$J&*&%"&1$*$76F$D67L>$@E$@-.OG-6.B-7D$+.:78;$38**H$32'H$
@&BH$SHOH$V2)#:'H$9%:'H6$M&)B"%$W1$WX2X';
Lobreau-Callen, D. 0%#/;$`:,:48c,6>$4@C+:,c>$BL$+@..67$B6>$<4:4-7:O
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C:8:46:6;$32'H$e&,)BH$75:'H$0/"1$"X02^0';
Louis, J. & Léonard, J. 0%^#; [48@576C:46:6;$H+;$'##2'%"$-71$V*2)#$
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Maas, P.J.M., Baas, P., Boesewinkel, F.D., Hiepko, P., Lobreau-
Callen, D., van den Oever, L. & ter Welle, B.J.H. 0%%';$K=6$
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Maheshwari, P., Johri, B.M. & Dixit, S.N. 0%!X;$ K =6$ E . @,: .$ C@ ,O
+=@.@DG$:7B$ 6C?,G@.@DG$@E$8=6$U@,:78=@-B6:6$ &U@,:78=:46:6(;$
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Maheshwari, P. & Singh, B. 0%!';$YC?, G@.@D G$@E$J&+)2:2*#%$+2+,"%G
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Malécot, V. & Nickrent, D.L. '//#;$T@.64L.:,$+=G.@D6768-4$,6.:8-@7O
>=-+>$@E$[.:4:46:6$:7B$,6.:86B$Q:78:.:.6>;$75:'H$32'H$""1$%X20/W;
Malécot, V., Nickrent, D.L., Baas, P., van den Oever, L. & Lobreau-
Callen, D. '//^;$*$C@,+=@.@D-4:.$4.:B->8-4$:7:.G>->$@E$[.:4:46:6;$
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Mathiasen, R.L., Nickrent , D.L., Shaw, D.C. & Watson, D.M. '//#;$
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McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, V., Hawksworth,
D.L., Marhold, K., Nicolson, D.H., Prado, J., Silva, P.C., Skog,
J.E., Wiersema, J.H. & Turland, N.J. (eds.). '//W;$9%'#)%&'"2%&*$
+2C#$2.$B2'&%"+&*$%20#%+*&'8)#$Xd"#%%&$A2C#Zg$IC21'#C$B5$',#$
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Nickrent, D.L. '//';$T->8.68@6$+=G.@D6768-4>1$ `L, ,678$ ,6.:8-@7>=-+>$
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Nickrent, D.L., Duff, R.J., Colwell, A.E., Wolfe, A.D., Young, N.D.,
Steiner, K.E. & dePamphilis, C.W. 0%%#;$T@.64L.: ,$+= G.@ D6768-4$
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Nickrent, D.L. & García, M.A. '//%;$[7$8=6$?,-7 5$@E$=@.@+:,:>-8->C1$
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Nickrent, D.L. & Malécot, V. '//0;$*$C@.64L.:,$+=G.@D67G$@E$Q:78:O
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4#)"%&1$KF@$76F$:,?@,6>4678$ D676,:$ @E$Q:78:.:46:6$E,@C$T:B:O
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TA XO N$!%$&'($)$*+,-.$'/0/1$!"#2!!#3-45,678$9$:.;$)$Classif ication of Santalales
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Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider,
H. & Wolf, P.G. '//W;$*$4.:>>-E-4:8-@7$ E@,$ 6S8:78$E6,7>;$ P& _ 2 %$
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,6.:86B$E@,C>;$N)#42%$7'&'#$J2%24)H$7'8CH$32'H$!1$02%";
Soltis, D.E., Clayton, J.W., Davis, C.C., Gitzendanner, M.A., Cheek,
M., Savolainen, V., Amorim, A.M. & Soltis, P.S. '//X;$T@7@O
+=G.G$:7B$ ,6.:8-@7>=-+>$ @E$8=6$ 67-DC:8-4$ :C+=-8,@+-4:.$E:C-.G$
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Soltis, D.E., Senters, A.E., Zanis, M.J., Kim, S., Thompson, J.D.,
Soltis, P. S., Ronse De Craene, L.P., Endress, P.K. & Farris, J.S.
'//";$IL776,:.6>$:,6$>->86,$8@$@8=6,$4@,6$6LB-4@8>1$<C+.-4:8-@7>$E@,$
8=6$6A@.L8-@7$@E$+678:C6,G;$I0#)H$eH$32'H$%/1$^W02^X/;
Soltis, D.E., Soltis, P.S., Chase, M.W., Mort, M.E., Albach, D.C., Za-
nis, M., Savolainen, V., Hahn, W.H., Hoot, S.B., Fay, M.F., Ax-
tell, M., Swensen, S.M., Prance, L.M., Kress, W.J., Nixon, K.C.
& Farris, J.S. '///;$ *7D-@>+6,C$+=G.@D67G$-7E6,,6B$ E,@C$ 0#Q$
,M3*J$)B+@J$:7B$&'13$>6NL6746>;$32'H$eH$@"%%H$72+H$0""1$"#02^W0;
Spitzer, V., Bordignon, S., Schenkel, E. & Marx, F. 0%%^;$<B678-E-4:O
8-@7$@E$%$:468G.67-4$E:88GO:4-B>J$%O=GB,@SG>86:,-4$:4-B$:7B$%J0/O6+O
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&Q:78:.:46:6(;$eH$I0#)H$N"*$A,#0H$72+H$X01$0"^"20"^#;
Spitzer, V., Tomberg, W., Hartmann, R. & Aichholz, R. 0%%X;$* 7: .GO
>->$@E$8=6$>66B$@-.$@E$M#":'#)"&$:"*<&%""$&[.:4:46:6(1$*$,-4=$>@L,46$@E$
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Stauffer, H.U. 0%W0:;$V6-8,}D6$ RLC$V.v867B-:D ,:CC$B6,$Q:78:.:.6>;$
d#),H$7+,c#"/H$S&'8).H$L#:;$0^01$0'"20'!;
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Stevens, P.F. '//W;$U;*;Q;$g@=7>@7$,6A-6F$7@;$!;$*7$67B$8@$:..$8=-7D>•$
H.:78>$:7B$8=6-,$7:C6>;$I8:')&*H$75:'H$32'H$0%1$00!20"";
Su, H.-J. & Hu, J.-M. '//#;$H=G.@D6768-4$,6.:8-@7>=-+>$@E$V:.:7@+=@O
,:46:6$:7B$Q:78:.:.6>$?:>6B$@7$E.@,:.$V$=@C6@8-4$D676>;$*?>8,:48$
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Thompson, W.A. 0%^%;$*$7:8L,:.$=G?,-B$@E$@2)&%',8:$0"+)&%',8:$:7B$
P81#"&$&%'&)+'"+&;$P)&%:H$i$-)2+H$K25H$72+H$S#c$j#&*&%C$XX1$'/#;
Thorne, R.F. 0%#";$H,@+@>6B$76F$,6:.-D7C678>$-7$8=6$*7D-@>+6, C:6;$
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Tieghem, P.v. 0#%!:;$QL,$.6$D,@L+6C678$B6>$6>+{46>$67$D67,6>$B:7>$.:$
8,-?L$B6>$I:-:B67B ,c6>$B6$.:$E:C-..6$B6>$U@,:78=:4c6>;$38**H$72+H$
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Tieghem, P.v. 0#%!?;$QL,$.6$D,@L+6C678$B6>$6>+{46>$67$D67 ,6>$B:7>$.:$
8,-?L$B6>$H>-88:4:78=c>$B6$.:$E:C-..6$B6>$U@,:78=:4c6>;$38**H$72+H$
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Tieghem, P.v. 0#%W ;$QL ,$.6 > $+= : 7c,@ D:C 6>$€ $@AL. 6$>: 7 >$7L 46. .6J $E@ , C :78 $
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Tieghem, P.v. 0#%X;$QL,$.6>$-7>6C-7c6>$€$@AL.6>$>:7>$7L46..6$E@,C:78$
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Tieghem, P.v. 0#%%:;$M6LS$D67,6>$7@LA6:LS$+@L,$.:$E:C-..6$B6>$`@LO
.:4c6>;$38**H$J8:H$M":'H$S&'H$X-&)":Z$!1$%X20//;
Tieghem, P.v. 0# %% ?;$ Q L, $ .6 >$ `@ L. :4 c6 >; $ I%%H$7+"H$S&'H6$32'HJ$Q6,;$#J$
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Tieghem, P.v. 0%/!;$QL,$ .6$ D67,6$[48@47{C6$4@7>-Bc,c$ 4@CC6$8G+6$
BsL76$E:C-..6$B->8-7486J$.6>$[48@47cC:4c6>;$eH$32'H$0%1$^!2!#;
Ulloa Ulloa, C., Nickrent, D.L., Whitefoord, C., & Kelly, D.L. $<7$
+,6>>;$M2%C8)2C#%C)2%J$:$76F$C@7@8G+-4$D67L>$@E$*+8:7B,:46:6$
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Venkata Rao, C. 0%W^;$[7$8=6$C@,+=@.@DG$@E$8=6$4:.G4L.L>;$eH$9%C"&%$
32'H$72+H$^'1W0#2W'#;
Vidal-Russell, R. & Nickrent, D.L. '//X;$*$ C@.64L.:,$+=G.@D67G$
@E$8=6$E6:8=6,G$C->8.68@6$J":2C#%C)80;$75:'H$32'H$"'1$!W/2!W#;
Vidal-Russell, R. & Nickrent, D.L. '//#:;$YA@.L8-@7:,G$,6.:8-@7>=-+>$
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