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ISSN1346-7565 ActaPhytotax.Geobet.63(1)4:1-49 (2012)
Chromosome Countsfor theAsianMyristicaceae(Magnoliales)
KAZuO OG[NuMAi,SHAwN K, Y.LuM2 AND HIRosHI ToBE3･'
iFaculi.v, ofHuman Lij? ]and Environtnental S('ience ,University ilf'Kochi, Kochi 780-8515, Jopan;
]'Natttrat Science& SeienceEducation,?L'JEN,b,v,ang7lechnologicalUhiversity,Singapore639798;
SGraduate Schovl ofScien Kcyeo,t Uoniversit ?y<,yot 6o06-8502 ,Jtu)an.
"[email protected],yoto-tt,acJP(inuthorforcorre,spondencel
Myristicacea ae f,amil ywith pantropic dailstribut iaondn sLster to the rest of the Magno] iale sar,e poorly
understood with respect to chromosome numbers. Here we report somatic chromosome numbers of [4
species in fiv eof the six Asian genera. Chromosome numbers were consistent within individu aglenera
but markedly diffe ramong the genera. Bascd on our observations, we concluded that the generie base
number is x = 20 in Endocomia, x = 22 in GL,mnacranthera and Knema, x = 25 in IL(vrist iacnda, x = 26
in Horsfietdia .AIthough reliable data on chrernosomc numbers are currently limite tdo Asian genera ,a
grea tdivergence in base numbers i sa marked contrast to the low melecular divergenee within the family,
casting a doubt to datin gMyristicaceae to be young,
Key words: chromosome number, Magnoliales, Myristicaceae
Myristicace aceomprise around 400 species Gronov. )T.welve (60% )of the 20 genera remain
of trees in about 20 genera more or les sequally chromosomally unknown (se e[[lab le1) .Even
distribute din the tropics worldwide: eight genera those reported are inconsiste nast ,in case of ilcly-
in Africa, six genera in Arnerica a,nd six genera ristica (se eTable 1).
in Southeas tAsia (d eWilde 20eO, see Table 1). In this paper we report the results ofan inves-
Molecular analyses show that Myristicaceae are tigation of somatic chromosome numbers of 14
placed in Magnoliales and are sister to the rest of species of five Southeast Asian genera (Encloco-
the order (Sauqu eett aL 2003, Steven s2001 on- mia W,J. de Wilde, Gymnacranth eHronas,fietd-
wards). Severa lmorphological characters, such ia, Knema, and ?Llyrist iicna M)yristicaceae ,and
as androecium and po]le nmorphology, were re- provide a brief note on chromosome counts previ-
cently investigate din Myristicaceae (Sauquetously reported. The chromosome counts, which
2003, Sauquet & Le Thomas 2003) ,but many we present fbr all the Asian genera except Para-
other morphological characters are yet poorly un- no,ristica W. J. de Wilde (P .sqpicana (Foreman)
derstood .In particula rch,romosome numbers W, J .de Wilde only), which is endemic to Papua
which often refiect evolutionary relationships New Guinea, provid ea better understanding of
among familie s(e.g .O,ginuma & Tbbe 2006: chromosome feature osfthe family.
Laurales) ,or within families (e. gO.g,inuma &
Tbbe 1997: Rhizophoracee), have been poorly Materials and Methods
studied in the famil y(fo rreview see KUhn & Ku-
'fable
bitzk i1993) .Chromosome counts publishe dso The 14 species examined are liste din 2
far are restricted to eight genera includin gfour along with their respective collection data and
Southeas tAsian genera (Clymnacranth Wearrab,, chrornosome numbers. Somatic chromosomes
Hbnvieldi aWilld. ,Knema Lour, and Mlyristica were examined using young leave scollected
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TARt.F .]. Summary efchromoseme informati eonn Myristicacea eand generjc base number. Chromoseme numbers reported
in prescnt study are indicate din bo]d.
Taxon Distribution Chromosomcnumbcrs Basenumber
Brochoneura Warb. AfricaAfricaAt'rica?Am?e?rica
Cephalosphaera Warb.
Coelocat'vonWarb,
Compsoneura(DC,)Warb. 2n=ca.102b
Dialyanther Waarb. AmericaAsiaAsiaAfrile?aAsiaAmerica
Endocomia 2n=402n=44 -.20x=22
x
G},mnacranthera
n=21?n=25
HinematodendhonCapuron
Hbnlfieldia 2n 52
=SO, x=26
Iu'anthera Warb. 2n = ]ooi'
Knemaimttioutchia AsiaAfricaAs]aAmerinc=a21A?mn=ericaAsiaAfrica2Anf-r4i4caAfricaAmericax-22
Warb.
oprLs'tica 19, 21, 26 2n=502n x=25
Osteophtoeum Warb. = ca. 2soM
Otoba(DC.)H.Karst. ??
Paramvrisitica
Pvcnanthi tWsarb. 2n == 3s4'
Sq}lphocephalittmWarb. ??
Sta!tdtWiaarb.
varoia Aubl. 2n = 52'J
Reference sfor previou schromoseme cou nts for species ef G.ymnact'antl] eHroan.vfieieS Ktnae,ma, and M.i,ristic aare presented in
text.h
Data from Compsoneura ulei Warb, (Morawet z1986}.
!) Data from j}lvanthera coriacea Ducke (Morawetz 1986).
i' Data from Osteophtoeumpiat.vsperm eWtamrb. (Morawet z1986).
J lData from Pyenanthus angoiens is (We[w. W}arb. CMangcnot & Mangenot 19S7).
"' Data from Yirot acatoph.J,lta Warb., J{ divergeny Ducke and U eiongala Warb. (Morawetz 1986).
,
from trees in the fiel dM.ethods of pretreatment, (King )W.J, de Wilde .The species had 2n - 40
fixatio nan,d staining fbr chromosomes fbllowed (Fig s1., 2). The chromosomes at metaphase grad-
Oginuma et aL (1992 )At. leas tthree to five cells ually varied in length from O.7 to 1. 7"m. Ofthe
were examined to determi ntehe chromosome 40 chromosomes, some had a centromere in a
number. median position ,but the centromere positio nin
Terminology of chromosome morphelogy on others was uncertain. The base number ofthis ge-
the basj softhe positio onfthe centromere foIlows nusisx=20.
Levan (1964).
et aL
Clymnacranthera
Results Discussion
and The genus G.vmnacranthera comprises seven
species distribut efdrem southern Peninsular
Chromosome counts ofindivi gcenheiraal Thailan dto New Guinea (de Wilde 2000). We
Endocomia studied two species G, farquhari aWanrab. var.
The genus Endocomia comprises four species fonquhari a(nFaig s3,. 4) ,and G. forbe sWiairb.
distribute dfi:om southern China to New Guinea (Fig s5., 6), both of which had 2n = 44. The chro-
(d eWilde 2000) .We observed chromosomes for mosomes at metaphase gradua]ly varied in length
the firs ttime for the genus using E. canarioides from O,3 to O.7 ptm. In both species, many of the
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1/ABLE 2. Taxa of Myristicaeea estudied, voucher inibrmatie nand chromosome numbcrs.
Taxon CoJlection Chromosome
number
Endocomia eananioides SINGAPORE.BukitTimah 2n=40
Oginuma & Lu,n 0701 (KYO)
G.ymnaerantherafarquhariana SINGAPORE, BukitTimah 2n=44
var.jurguhaniana Oginuma & Lttm 0702 (KYO)
G.forbesii SINGAPORE.MacRitchie 2n=44
Oginuma & Lum 9703 (KYO)
Hbi'sheldiaitl}'a Cu]tivateSdi,ngaporeBotanicaGlarden. 2n! 52
Oginuma & Lum 0704 (KYO)
ll. S]NGAPORE.MacRitchie 2n=52
sucosa
Qginttt &n aLitm 0705 (KYO)
H. CultivatcdS,ingaporeBotanicaG]arden. 2n-52
si{perba
Oginuma & Lutn 0706 (KYO)
H, walliehii SINGAPORE. Bukit Timah 2n=52
Ogint,ma& Lztm0707(KYO)
Knemajut:titracea SINGAPORE. BukitTimah 2n 44
=
Oginunia& Lun?0708(KYO)
K. gtobutaria SINGAPORE. Ubin Is, 2n=44
Oginuma & Lum 0709(KYO)
K, latericicr Cultivated,Singapore BotanicalGarden. 2n-44
Oginuma & Lum 0710 (KYO)
K.taurina Cultivated,SingaporeBotanicaGlarden. 2n-44
Oginuma & Lu,n 07Jl (KYO)
Mvristicacinnamoniea SINGAPORE.MacRitchie, 2n=50
Oginuma & Lum 07i2 (KYO)
M eUiptica Cultivated ,Singapore BotanicaL Garden, 2n = 5e
Oginuma & Lum 0713 (KYO)
Mfragrans Cultivated.SingaporeBotanicatGarden. 2n=5e
Oginttma & Lum 07i4 (KYO)
44 chromosomes had a centromere in a median Hbrshetdia
positio nb,ut the centromere positio nin others The genus Honxf7eldi acomprises about 100
was uncertaln. species that occur from Sri Lanka and southern
There are three previous records on chromo- China east to the Solomon Islands and northern
somc counts of Gymnacranthera :n = 2] from G. Australia (d eWilde 2000) .We studied fbur spe-
forbe s(iKiUhn & Kubitzki 1993) and G, panicu- cies H. irva Warb., H. sucosa Warb, (Fig s7., 8),
lat aWarb. (Ehrendor eft eaLr 1968) ,and 2n ; 44 Hl smperha Warb. (Fig s9, .10) ,and H. wallichii
fi'om G. paniculata (Oginum aet aL 1999 )(Tab]e Warb. (Fig s11., 12) ,which all had 2n - 52, Chro-
1) ,The chromosome number of2n = 44 fi'o mthe mosomes at metaphase of the three species grad-
three species examined (i. eG.. .,farquhariana, G. ually varied in length from O.3 to 1.0 ltm .Of the
forbes ianid, G, panicuiata) indicat etshat the 52 chromosomes, rnany had a centromere in a
base number of dytnnacrant hise xr =a 22. The median positio nb,ut the centromere positio inn
chromosome ceunt n = 21 reported for G. forbesiiethers was uncertain
and G. panicutata appears to be erroneous, at The previous chromosome counts fbr Hors-
leas trequiring confirmation. .fietdi aare n = 25 (Ehrendor etf ealr. 1968) and 2n
= 50 (Morawet z1986) fbr H. oclorata Willd .The
four species examined in our study (i. eH.. ,iilva,
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∴
麟 轟
．
3
べ』彗
鴪 轄 ． ・・
舜 泌 3
隙 轡
巒
愚
螺
灣
Fiα 1−6． Somatic chromosomes at mitotic metaphase in Myristicaceae ．1，3，5：micrographs ．2，4，6：drawings ofmicrographs
in l，3，and 5， respectively 」1，2：Endoco〃軍’αcanarioides （2n＝40）．3，4：Gymnaeranthera丿earguhar ’ana var． farquhイaanaJ
（2月＝44）．5，6：G．ノbrbesi’（2n＝44＞Scal ebar＝2μm ．
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定
喚
●
・蝕 ・鞭’炉
〆
●
ψ
秘塾
ぜ
〆
弄
騨
メ
∵
’
∴ 駕 宀 ． 馬．’ 7 ．
る ・， ぎ 塙 レ 「・ 8 し
読∴ ゴ● の磁 飢 ジ 6 ●ゆ 餮 ■
捻駕聴 曜〜 Q ・、ゆ 鷲 噸
・
ぐ
即華 櫑 ：
鑼 鱒汐 静
・
魯●◆R
FIG，7−12， Somatic chromosomes at mitotic metaphase in Myristicaceae．7，9， l l：micrographs ．8，10，12：drawings of micro −
g呂r5a2p）h．s Sicna 7l，e b9a， ran＝d 2l μl，m re．spective 【y．7，8：Horsfi’ゴe’αsucesa （2η＝52）．9，10：H． supe 厂わo （2n＝52）．1亘，12：H． wallichii （2n
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H sukosa, U superba, and H waUichii) had 2n = zaylanica Thwaites (Morawe t1z986) .Our counts
52, indicatin ga base number ofx = 26 in Hbnst7el- of three species agreed only with Morawetz
dia .The previou schromosome counts of n = 25 (1986 C)h.romosome numbers may vary within
and 2n = 50 for IL odorata appears to be incoF the genus ,but our results indicat tehat the base
rect and require confirmation. number of torisit iis cx a= 25.
Khema Chromosome inILlyristicaceae
evolution
The genus Khema comprises about 90 species In the presen tstudy we determine dthe chro-
that oocur from India to western New Guinea (de mosome base number for the fiv eAsian genera of
Wilde 2000) .We studied four species of Khema: Myristicaceae :x = 20 in Entlocomia, x = 22 in
K fiti:liiWraarbc.e,aK. Warb.(Figs,(lymnacranthera Khema, 25in?l(lyristi-
globularia and x =
13, 14), KL laterici aElmer (Fig s15., 16) ,and KL ca, and x = 26 in Hbnstfeldia .It is premature to
laurin aWarb. (Fig s17. ,18) ,which all had 2n = discus schromosome evolution within the family
44. Chromosomes at metaphase of the four spe- becaus eexact chromosome informati oonn My-
cies gradually varied in lengt hfrom O.3 to 1.2 risticaceae is restricted to these fiye Asian gen-
pam .Some of the 44 chromosornes had a centro- era. Ofinteres th,owever ,is the fac tthat the Lau-
mere in a median, submedian, or subterminal po- rales, which are considered to be sister to the
sition, but the centromere positio inn others was Magnoliale s(e, gS,o,lti set al, 2011) ,have x = 11
uncertam. as a probabl earchaic base number (Oginum &a
Ehrendorfe ret al. (196 8p)reviousl ryeported Tbbe 2006) .Ifx= 11 is plesiomorphi icn Magno--
a count for KL intermedia (Blum eW)arb. of n = liale ass well, the base number x = 22 (i n([lymna-
21. All four species examined in our study (K, eranthera and Khema) might have been derived
juderace aK, globularia ,K laterici aan,d K. by polyploidizat ifornom x = 11. The other num-
Iaurina )had 2n = 44, indicati ntghat the base bers x = 20, 25, and 26 may have been derive dby
number of Khema is x = 22. The ehromosome an aneuploid decreas eand increas efrom x = 22.
count n = 21 of KL intermedia requires confirma- Compared to other farnili oefs the Magnolia-
tion. lesA,nnonaceaetr 7,8,9) Magnoliaceaeix
= and
= 19) (Okad a1975) ,Myristicaceae have a great
dyristica diversi tiyn generi cbase numbers, with at leas tx
The genus ILdyristi ccaomprises about l70 = 20, 22, 25, 26. In contrast, Doyle et al. (2004)
species that occur from India to northern Austra- showed strikingly low molecular divergence,
li aand far int othe Pacifi c(d eWilde 2000) .We based on several plasti adnalyses, in the Myristi-
studied three species of ?Llyristi c?at:d lcinnamo- caeae. An estimated age by molecular dating for
mea King (Fig s19,, 20) ,Ml elliptica Wall. (Figs.the crown-group ofthe family was 15-18 or 17-
21, 22) ,and IL d.[fragrans Houtt .(Fig 2s3., 24) ,All 21 Myr, suggesting either a young age for the
had 2n = 50. Chromosomes at metaphase gradu- family or a marlced slowdown in molecular evolu-
ally varied in length from O.3 to 1.2 ptm. In all the tion (Doyl eet aL 2004) .Doyle et al. (2004)'fuF
three species examined, some of the 50 chromo- ther discus sthat the oldest diagnost ifcossi losf
somes had a centromere in a median, submedian, the Myristicaceae are Miocene seeds which can
or subterminal positio nb,ut the centromere posi- be taken as evidence for the young age of the
tion in others was uncertain. family ,but tha tis implausib lfebr the transoce-
Various chromosome numbers have been re- ani¢ dispers alof the large ,animal-dispersed
corded preyiously :n = 19 in Ml insipidu R. Br. seeds in the family A.lthough exact chromosome
(Sol t&i sSolti s1990) ,n - 21 in Ml elliptica (Eh- data are currently limited to the Asian taxa, a
rendorfer et al. 1968) and M .fragrans (Simmondsgrea tdiversi tiyn their bas enumbers casts doubt
1954), n = 26 in M erratica Hook. f & Thomp- on a young date for Myristicacea eb,ut instead
son (Mehr a& Bawa 1969), and 2n = 50 from ?Ld: suggests that the family is much older. Further
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売・腰榊瓜’ー
欝 纏び 5 鍾默簸丶
鞭 響
6 。噛
ミ 98 D
騨 ●
蘇馨 の 、 。 （ も ら
調 華 偽ε 、3♂ ．・ 亀 ・． ・ ● ● ●
鱒 騨 繕 硫 等 ・ 鵬
解 ：、 濕 te ボ吟 漕譖
禪 ぴダ 「。 雄醗
r・丶 燐 艶
ザ 舮 3、84 燭
梅 も
噂 雪
亀 、
灘 ．
．瞭 帽
譁
ρ ゜
宀
聯ド
も θ 9
噸
3
∂ ■ 噸 ◎ ρ 塾
3
∴
含
・
、 ，
“こ勢 、 、
．
8
％ 鳩 ご
FIG．13−18． Somatic chromosomesat mitoIic metaphase in Myristicaceae ．13，15，17：micrographs ．14，16，18：drawings ofmi −
crographs in 13，亘5， and 17， respectively ．13，14：Kne〃軍αg’obularia （2n；44）．15，艮6：K． laterici（a2n；44）．17，18：K． laπ一
r加α（2n≡44）． Scal ebar‘2μm ．
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瀞劇
灘
転
凶
輔雛
澤
濘
ビ
’
’
亀
帆 ・・擁 弓 「
夛
鬱
3
三
歌 築
！
30
り ゜φ ，・ ●
er
あ
・ ・ ’
● σ
凭
瀦
FIG．19−24． Somatic chronlosomes at mitotic metaphase in Myristicaceae．19，21，23：micrographs ．20，22，24：drawing sofmi −
ノc如roggrrap側hs∫（ 2inn ≡195，02）1． ， Sacnadl 2e b3a， rre己spe2ctμivmel．y ．19，20：Myristi ccainamonea （2n；50）．21，22：M ．θ〃iptica（2n＝50）．23，24：M ．
一
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studies are needed of chromosome numbers in G.vnotroch easn,d Pettaca4y ('Rxhizophorac eGaye-;
the African and American taxa. notrocheae). Aeta Phytotax. Geobot .48: 15-21.
OgLi.numa, K. & H. Tobe .2006. Chromosome cvolution in
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