<i>Quambalaria</i> species associated with eucalypt diseases in southern China

doi:10.15302/J-FASE-2017173

Front. Agr. Sci. Eng.

2017, Vol. 4

Issue (4) : 433-447 https://doi.org/10.15302/J-FASE-2017173

RESEARCH ARTICLE

Quambalaria species associated with eucalypt diseases in southern China

Shuaifei CHEN^1,², Qianli LIU¹, Guoqing LI¹, Michael J. WINGFIELD²(

)

¹. China Eucalypt Research Centre, Chinese Academy of Forestry, Zhanjiang 524022, China
². Forestry and Agricultural Biotechnology Institute, University of Pretoria, Pretoria 0028, South Africa

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Abstract

The genus Quambalaria includes several important pathogens of species of Eucalyptus and Corymbia, mainly causing leaf and shoot blight. Recently, extensive shoot and leaf dieback and stem cankers suspected to be Quambalaria diseases have been found on young Eucalyptus urophylla × E. grandis trees in Guangdong and Hainan Provinces. The occurrence of Quambalaria species and their association with eucalypt hosts within China needs to be investigated for tree diseases management. The isolates from the diseased samples were identified based on their morphological structures and phylogenetic analyses with DNA sequence data for the internal transcribed spacer region and large ribosome subunit RNA of the nuclear rDNA. This work revealed that three species of Quambalaria were present: Quambalaria pitereka from Corymbia citriodora, Q. eucalypti from E. urophylla × E. grandis, both isolated from young eucalypt leaves and shoots in Guangdong Province, and Quambalaria simpsonii, which was isolated from stem cankers of E. urophylla × E. grandis at four different sites across Guangdong and Hainan Provinces. These results confirmed that Quambalaria agents were associated with the diseases occurring on eucalypt hosts in South China. This is the first report of Q. eucalypti in Asia and the first report of Q. simpsonii in China on Eucalyptus trees.

Keywords Corymbia Eucalyptus forest pathogens plantations Myrtaceae

Corresponding Author(s): Michael J. WINGFIELD

Just Accepted Date: 30 October 2017 Online First Date: 22 November 2017 Issue Date: 10 December 2017

Cite this article:

Shuaifei CHEN,Qianli LIU,Guoqing LI, et al. Quambalaria species associated with eucalypt diseases in southern China[J]. Front. Agr. Sci. Eng. , 2017, 4(4): 433-447.

URL:

https://academic.hep.com.cn/fase/EN/10.15302/J-FASE-2017173
https://academic.hep.com.cn/fase/EN/Y2017/V4/I4/433

Fig.1 Symptoms of infection by Quambalaria spp. on eucalypt trees. Shoot (a) and juvenile leave (b) of Corymbia citriodora infected by Quambalaria pitereka covered in white masses of conidia and conidiophores. New shoot (c) produced from the infected C. citriodora, and reinfected by Q. pitereka. Death of apical shoot (d) of Eucalyptus urophylla× E. grandis clone infected by Quambalaria simpsonii. Mature leaf (e) and young apical shoot (f) of E. urophylla× E. grandis clone infected by Q. simpsonii. Arrows indicate infected sites.

Identity	Isolate No.^a	GenBank accession No.^b		Host	Location	Collector	Reference
Identity	Isolate No.^a	ITS	LSU	Host	Location	Collector	Reference
Quambalaria coyrecup	WAC12947^cde	DQ823431	DQ823444	Corymbia calophylla	Western Australia, Australia	T Paap	Paap et al.^[23^]
Q. coyrecup	WAC12948^de	DQ823433	DQ823446	C. calophylla	Western Australia, Australia	T Paap	Paap et al.^[23^]
Q. coyrecup	WAC12949^e	DQ823432	DQ823445	C. calophylla	Western Australia, Australia	T Paap	Paap et al.^[23^]
Q. coyrecup	WAC12950^de	DQ823429	DQ823447	C. calophylla	Western Australia, Australia	T Paap	Paap et al.^[23^]
Q. coyrecup	WAC12951^de	DQ823430	DQ823448	C. calophylla	Western Australia, Australia	T Paap	Paap et al.^[23^]
Q. coyrecup	BRIP48338^d	EF444877	N/A^g	C. polycarpa	Darwin, Northern Territory, Australia	R Pitkethley	Pegg et al.^[24^]
Q. coyrecup	BRIP48339^d	EF444878	N/A	C. polycarpa	Darwin, Northern Territory, Australia	R Pitkethley	Pegg et al.^[24^]
Q. cyanescens	CBS357.73^cde = CMW5583	DQ317622	DQ317615	skin of man	Netherlands	TF Visser	de Beer et al.^[22^]
Q. cyanescens	CBS876.73^de = CMW5584	DQ317623	DQ317616	Eucalyptus pauciflora	New South Wales, Australia	VF Brown	de Beer et al.^[22^]
Q. cyanescens	WAC12952^de	DQ823419	DQ823440	C. calophylla	Western Australia, Australia	T Paap	Paap et al.^[23^]
Q. cyanescens	WAC12953^de	DQ823422	DQ823443	C. ficifolia	Western Australia, Australia	T Paap	Paap et al.^[23^]
Q. cyanescens	WAC12954^e	DQ823420	DQ823442	C. calophylla	Western Australia, Australia	T Paap	Paap et al.^[23^]
Q. cyanescens	WAC12955^de	DQ823421	DQ823441	C. calophylla	Western Australia, Australia	T Paap	Paap et al.^[23^]
Q. cyanescens	BRIP48396^d	EF444874	N/A	Native C. citriodora	Beaudesert, Queensland, Australia	GS Pegg	Pegg et al.^[24^]
Q. cyanescens	BRIP48398^d	EF444875	N/A	Native C. citriodora	Beaudesert, Queensland, Australia	GS Pegg	Pegg et al.^[24^]
Q. cyanescens	BRIP48403^d	EF444876	N/A	Native C. citriodora	Beaudesert, Queensland, Australia	GS Pegg	Pegg et al.^[24^]
Q. eucalypti	CBS118844^cde = CMW1101	DQ317625	DQ317618	Eucalyptus grandis	Kwambonambi, South Africa	MJ Wingfield	de Beer et al.^[22^]
Q. eucalypti	CBS 119680^de = CMW11678	DQ317626	DQ317619	E. grandis clone NH58	Kwambonambi, South Africa	L Lombard	de Beer et al.^[22^]
Q. eucalypti	CMW14329	DQ317614	N/A	E. grandis× E. camaldulensis clone	Kwambonambi, South Africa	J Roux	Roux et al.^[30^]
Q. eucalypti	CBS118615= CMW17252	DQ317609	N/A	E. nitens	Rooihoogte, South Africa	ZL Mthalane & J Roux	Roux et al.^[30^]
Q. eucalypti	CMW17253	DQ317610	N/A	E. nitens	Rooihoogte, South Africa	ZL Mthalane & J Roux	Roux et al.^[30^]
Q. eucalypti	CMW17254	DQ317611	N/A	E. nitens	Rooihoogte, South Africa	ZL Mthalane & J Roux	Roux et al.^[30^]
Q. eucalypti	CMW17255	DQ317612	N/A	E. nitens	Rooihoogte, South Africa	ZL Mthalane & J Roux	Roux et al.^[30^]
Q. eucalypti	CBS118616= CMW17771	DQ317613	N/A	E. grandis clone	Kwambonambi, South Africa	J Roux	Roux et al.^[30^]
Q. eucalypti	UY1036	EU439922	N/A	Myrceugenia glaucescens	Uruguay	C. A. Pérez	Pérez et al.^[28^]
Q. eucalypti	UY1718	EU439923	N/A	M. glaucescens	Uruguay	C. A. Pérez	Pérez et al.^[28^]
Q. eucalypti	PE3/MEAN 996	JX297605	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE6/MEAN 997	JX297603	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE27/MEAN 998	JX297604	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE28/MEAN 999	JX297600	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE29/MEAN 1000	JX297602	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE30/MEAN 1001	JX297601	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE52/MEAN 1002	JX297606	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE53/MEAN 1003	JX297598	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE54/MEAN 1004	JX297599	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE93/MEAN 1006	KR336802	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE96/MEAN 1009	KR336803	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE151/MEAN 1012	KR336804	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE152/MEAN 1013	KR336805	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE153/MEAN 1014	KR336806	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	PE154/MEAN 1015	KR336807	N/A	E. globulus	Portugal	N/A	Braganca et al.^[27^]
Q. eucalypti	BRIP48367	EF444823	N/A	C. torelliana× C. citriodora subsp. variegata	Walkamin, Queensland, Australia	GS Pegg	Pegg et al.^[24^]
Q. eucalypti	BRIP48422^d	EF444832	N/A	E. dunnii	New South Wales, Australia	AJ Carnegie	Pegg et al.^[24^]
Q. eucalypti	BRIP48498^d	EF444844	N/A	E. grandis	New South Wales, Australia	AJ Carnegie	Pegg et al.^[24^]
Q. eucalypti	BRIP48507^d	EF444822	N/A	E. grandis	Moggill, Queensland, Australia	GS Pegg	Pegg et al.^[24^]
Q. eucalypti	CERC8476^d	KY615009	N/A	E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. eucalypti	CERC8477^g	KY615010	N/A	E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. eucalypti	CERC8478	KY615011	N/A	E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. eucalypti	CERC8479^g	KY615012	KY615050	E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. eucalypti	CERC8480^g	KY615013	N/A	E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. eucalypti	CERC8481	KY615014	KY615051	E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. eucalypti	CERC8482^g	KY615015	N/A	E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. eucalypti	CERC8483	KY615016	N/A	E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. pitereka	DAR19773^cde	DQ823423	DQ823438	C. eximia	New South Wales, Australia	J Walker & AL Bertus	Paap et al.^[23^]
Q. pitereka	CMW 6707^de	DQ317627	DQ317620	Corymbia maculata	New South Wales, Australia	MJ Wingfield	de Beer et al.^[22^]
Q. pitereka	CBS118828^de = CMW5318	DQ317628	DQ317621	C. citriodora subsp. variegata	Queensland, Australia	M Ivory	de Beer et al.^[22^]
Q. pitereka	CMW23610	EF427372	N/A	C. citriodora	Guangdong, China	YJ Xie	Zhou et al.^[17^]
Q. pitereka	CMW23611	EF427373	N/A	C. citriodora	Guangdong, China	YJ Xie	Zhou et al.^[17^]
Q. pitereka	CMW23612	EF427374	N/A	C. citriodora	Guangdong, China	YJ Xie	Zhou et al.^[17^]
Q. pitereka	CMW23613^d	EF427375	N/A	C. citriodora	Guangdong, China	YJ Xie	Zhou et al.^[17^]
Q. pitereka	BRIP48325	EF427366	N/A	C. citriodora subsp. variegata	Queensland, Australia	GS Pegg	Zhou et al.^[17^]
Q. pitereka	BRIP48361^d	EF427367	N/A	C. citriodora subsp. variegata	Queensland, Australia	GS Pegg	Zhou et al.^[17^]
Q. pitereka	BRIP48370^d	EF427368	N/A	C. citriodora subsp. variegata	Queensland, Australia	GS Pegg	Zhou et al.^[17^]
Q. pitereka	BRIP48384^d	EF427369	N/A	C. citriodora subsp. variegata	Queensland, Australia	GS Pegg	Zhou et al.^[17^]
Q. pitereka	BRIP48386^ed	EF427370	N/A	C. citriodora subsp. variegata	Queensland, Australia	GS Pegg	Zhou et al.^[17^]
Q. pitereka	BRIP48531^d	EF427371	N/A	C. citriodora subsp. variegata	Queensland, Australia	GS Pegg	Zhou et al.^[17^]
Q. pitereka	WAC12957^e	DQ823426	DQ823437	C. ficifolia	Western Australia, Australia	T Paap	Paap et al.^[23^]
Q. pitereka	WAC12958^e	DQ823427	DQ823436	C. calophylla	Western Australia, Australia	T Paap	Paap et al.^[23^]
Q. pitereka	QP26^e	DQ823424	DQ823434	C. citriodora subsp. variegata	Queensland, Australia	GS Pegg	Paap et al.^[23^]
Q. pitereka	QP45^de	DQ823425	DQ823439	C. citriodora subsp. variegata	Queensland, Australia	GS Pegg	Paap et al.^[23^]
Q. pitereka	BRIP48346^d	EF444845	N/A	C. citriodora subsp. citriodora	Davies Creek, Queensland, Australia	GS Pegg	Pegg et al.^[24^]
Q. pitereka	BRIP48317	EF444854	N/A	C. henryi	Coolabunia, Queensland, Australia	GS Pegg	Pegg et al.^[24^]
Q. pitereka	BRIP48381^d	EF444858	N/A	C. citriodora subsp. citriodora	Silkwood, Queensland, Australia	GS Pegg	Pegg et al.^[24^]
Q. pitereka	BRIP48383^d	EF444859	N/A	C. citriodora subsp. variegata	Beaudesert, Queensland, Australia	GS Pegg	Pegg et al.^[24^]
Q. pitereka	WAC12956^d	DQ823428	N/A	C. ficifolia	Western Australia, Australia	T Paap	Paap et al.^[23^], Pegg et al.^[24^]
Q. pitereka	BRIP48349^d	EF444860	N/A	C. torelliana × C. citriodora subsp. variegata	Mareeba, Queensland, Australia	GS Pegg	Pegg et al.^[24^]
Q. pitereka	BRIP48325^d	EF427366	N/A	C. citriodora subsp. variegata	Binjour, Queensland, Australia	GS Pegg	Pegg et al.^[24^]
Q. pitereka	BRIP48328^d	EF444872	N/A	Native C. citriodora subsp. variegata	Dilkoon, New South Wales, Australia	GS Pegg	Pegg et al.^[24^]
Q. pitereka	BRIP48432^d	EF444873	N/A	C. citriodora subsp. variegata	Grafton, New South Wales, Australia	GS Pegg	Pegg et al.^[24^]
Q. pitereka	CERC8486^de	KY615017	KY615052	C. citriodora provenance CERC10	Guangdong, China	SF Chen & GQ Li	This study
Q. pitereka	CERC8488^e	KY615018	KY615053	C. citriodora provenance CERC12	Guangdong, China	SF Chen & GQ Li	This study
Q. pitereka	CERC8489	KY615019	N/A	C. citriodora provenance CERC13	Guangdong, China	SF Chen & GQ Li	This study
Q. pitereka	CERC8491	KY615020	N/A	C. citriodora provenance CERC15	Guangdong, China	SF Chen & GQ Li	This study
Q. pitereka	CERC8494^eg	KY615021	KY615054	C. citriodora provenance CERC17	Guangdong, China	SF Chen & GQ Li	This study
Q. pitereka	CERC9093	KY615022	N/A	C. citriodora provenance CR76	Guangdong, China	SF Chen & Y Lin	This study
Q. pitereka	CERC9094	KY615023	N/A	C. citriodora provenance N371	Guangdong, China	SF Chen & Y Lin	This study
Q. pitereka	CERC9095	KY615024	N/A	C. citriodora provenance N28	Guangdong, China	SF Chen & Y Lin	This study
Q. pitereka	CERC9096	KY615025	N/A	C. citriodora provenance N411	Guangdong, China	SF Chen & Y Lin	This study
Q. pitereka	CERC9097^eg	KY615026	KY615055	C. citriodora provenance N223	Guangdong, China	SF Chen & Y Lin	This study
Q. pitereka	CERC9098^g	KY615027	N/A	C. citriodora provenance N322	Guangdong, China	SF Chen & Y Lin	This study
Q. pitereka	CERC9099^eg	KY615028	KY615056	C. citriodora provenance CR033	Guangdong, China	SF Chen & Y Lin	This study
Q. pitereka	CERC9100	KY615029	N/A	C. citriodora provenance CR039	Guangdong, China	SF Chen & Y Lin	This study
Q. pitereka	CERC9101	KY615030	N/A	C. citriodora provenance CR92	Guangdong, China	SF Chen & Y Lin	This study
Q. pitereka	CERC9102	KY615031	N/A	C. citriodora provenance CR36	Guangdong, China	SF Chen & Y Lin	This study
Q. pitereka	CERC9103^e	KY615032	KY615057	C. citriodora provenance N601	Guangdong, China	SF Chen & Y Lin	This study
Q. pitereka	CERC9104	KY615033	N/A	C. citriodora provenance N28	Guangdong, China	SF Chen & Y Lin	This study
Q. simpsonii	CBS 124772^de	GQ303290	GQ303321	Eucalyptus tintinnans	Edith Falls, Australia	BA Summerell	Cheewangkoon et al.^[26^]
Q. simpsonii	CBS 124773^de	GQ303291	GQ303322	Eucalyptus sp.	Lamphoon, Thailand	R Cheewangkoon	Cheewangkoon et al.^[26^]
Q. simpsonii	CERC8496^dg	KY615034	N/A	E. urophylla × E. grandis	Hainan, China	SF Chen & QL Liu	This study
Q. simpsonii	CERC8499	KY615035	N/A	E. urophylla × E. grandis	Hainan, China	SF Chen & QL Liu	This study
Q. simpsonii	CERC8505^d	KY615036	N/A	E. urophylla × E. grandis	Hainan, China	SF Chen & QL Liu	This study
Q. simpsonii	CERC8507^de	KY615037	KY615058	E. urophylla × E. grandis	Hainan, China	SF Chen & QL Liu	This study
Q. simpsonii	CERC8512^d	KY615038	N/A	E. urophylla × E. grandis	Hainan, China	SF Chen & QL Liu	This study
Q. simpsonii	CERC8514	KY615039	N/A	E. urophylla × E. grandis	Hainan, China	SF Chen & QL Liu	This study
Q. simpsonii	CERC8516	KY615040	N/A	E. urophylla × E. grandis	Hainan, China	SF Chen & QL Liu	This study
Q. simpsonii	CERC8517^e	KY615041	KY615059	E. urophylla × E. grandis	Hainan, China	SF Chen & QL Liu	This study
Q. simpsonii	CERC8519^dg	KY615042	N/A	E. urophylla × E. grandis	Hainan, China	SF Chen & QL Liu	This study
Q. simpsonii	CERC8526	KY615043	N/A	E. urophylla × E. grandis	Hainan, China	SF Chen & QL Liu	This study
Q. simpsonii	CERC8532	KY615044	N/A	E. urophylla × E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. simpsonii	CERC8534^deg	KY615045	KY615060	E. urophylla × E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. simpsonii	CERC8536^e	KY615046	KY615061	E. urophylla × E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. simpsonii	CERC8539^eg	KY615047	KY615062	E. urophylla × E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. simpsonii	CERC8541^d	KY615048	N/A	E. urophylla × E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Q. simpsonii	CERC8543^d	KY615049	N/A	E. urophylla × E. grandis	Guangdong, China	SF Chen & JQ Li	This study
Microstroma juglandis	R.B. 2042^de	DQ317634	DQ317617	Juglans regia	Germany	R Bauer	de Beer et al.^[22^]

Tab.1 Isolates of Quambalaria species collected from eucalypt trees in southern China in 2015 and 2016 and used for phylogenetic and morphological analysis

Fig.2 Phylogenetic tree based on maximum likelihood analysis of ITS sequence data of haplotypes of five Quambalaria species, Q. coyrecup (QCO), Q. cyanescens (QCY), Q. eucalypti (QE), Q. pitereka (QP) and Q. simpsonii (QS). Bootstrap values>60% are presented at branches, bootstrap values<60% or absent values are not shown. Haplotypes and isolates from eucalypts in this study are in boldface and highlighted. Isolates representing ex-type are marked with T, isolates repeated are marked with R. The tree is rooted to Microstroma juglandis.

Tab.2 Four haplotypes of Q. eucalypti as determined from the polymorphic nucleotides within the aligned sequence data of ITS region for isolates collected from species of Eucalyptus, C. torelliana× C. citriodora subsp. variegate and M. glaucescens

Tab.3 Twelve haplotypes of Q. pitereka as determined from the polymorphic nucleotides within the aligned sequence data of ITS region for isolates collected from species of Corymbia

Tab.4 Six haplotypes of Q. simpsonii as determined from the polymorphic nucleotides within the aligned sequence data of ITS region for isolates collected from species of Eucalyptus

Fig.3 Phylogenetic tree based on Maximum Likelihood analysis of large subunit sequence data of haplotypes of five Quambalaria species, Q. coyrecup (QCO), Q. cyanescens (QCY), Q. eucalypti (QE), Q. pitereka (QP) and Q. simpsonii (QS), respectively. Bootstrap values>60% are presented at branches, bootstrap values<60% or absent values are not shown. Haplotypes and isolates from eucalypts in this study are in boldface and highlighted. Isolates representing ex-type are marked with T, isolates repeated are marked with R. The tree is rooted to Microstroma juglandis.

Fig.4 Cultures grown on malt extract agar at 25°C after 2 weeks and the primary and secondary conidia. (a,b) Quambalaria pitereka; (c,d) Q. eucalypti; (e,f) Q. simpsonii.

Species	Isolate No.	Primary conidia			Secondary conidia
Species	Isolate No.	(L × W) size^a/mm	(L × W) mean^b/mm	L/W^c	(L × W) sizea/mm	(L × W) mean^b/mm	L/W^c
Q. pitereka	CERC8494	(7.0–)7.5–13.0(–20.5) × (2.5–)3.0–3.5(–4.0)	10.2 × 3.3	3.1	(4.0–)4.5–6.0(–7.0) × 2.0–2.5(–3.0)	5.2 × 2.5	2.1
	CERC9097	(8.0–)8.5–12.5(–15.0) × (2.5–)2.5–3.5(–4.0)	10.5 × 3.0	3.5	(4.5–)5.0–6.5(–7.0) × 2.0–2.5(–3.0)	5.7 × 2.3	2.5
	CERC9098	(8.0–)10.5–14.0(–16.0) × (2.0–)3.0–4.5(–5.0)	12.2 × 3.6	3.4	6.0–7.5(–8.0) × (2.0–)2.5–3.5(–4.0)	6.8 × 3.0	2.3
	CERC9099	(8.0–)9.0–12.5(–15.0) × (3.0–)3.5–4.5(–5.5)	10.7 × 4.0	2.7	5.5–7.0(–7.5) × 2.5–3.5(–4.0)	6.3 × 3.1	2.2
	Average^d	(7.0–)8.5–13.0(–20.5) × (2.0–)2.5–4.0(–5.5)	10.9 × 3.4	3.2	(4.0–)5.0–7.0(–78.0) × 2.0–3.0(–4.0)	6.0 × 2.7	2.2
Q. eucalypti	CERC8477	(4.5–)5.0–7.5(–8.0) × (3.0–)3.5–4.5(–4.5)	6.2 × 3.9	1.6	2.5–3.0(–3.5) × 2.5–3.0	3.0 × 2.5	1.2
	CERC8479	(5.5–)6.0–7.0(–7.5) × (2.5–)3.0–4.0(–4.5)	6.3 × 3.7	1.7	2.5–3.5(–4.0) × (2.0–)2.5–3.0	3.0 × 2.5	1.2
	CERC8482	(4.5–)5.0–6.5(–7.0) × (3.0–)3.5–4.0(–4.5)	5.7 × 3.9	1.5	3.0–4.0(–5.0) × 2.5–3.0	3.6 × 2.8	1.3
	CERC8480	(5.5–)6.0–7.0(–8.0) × (3.0–)3.5–4.0(–4.5)	6.4 × 3.8	1.7	3.0–3.6(–4.0) × 2.5–3.0	3.4 × 2.7	1.3
	Average^d	(4.5–)5.5–7.0(–8.0) × (2.5–)3.5–4.0(–4.5)	6.2 × 3.8	1.6	(2.5–)3.0–4.0(–5.0) × (2.0–)2.5–3.0	3.3 × 2.6	1.3
Q. simpsonii	CERC8496	(6.0–)6.5–9.5(–11.0) × (2.0–)2.5–3.5(–4.0)	8.3 × 3.1	2.7	(3.0–)3.5–4.5(–5.0) × 2.0–2.5(–3.0)	4.0 × 2.4	1.7
	CERC8519	(5.5–)6.0–8.0(–9.0) × 3.0–4.0(–4.5)	7.1 × 3.6	2.0	(2.5–)3.0–3.5(–4.0) × 2.0–2.5	3.3 × 2.3	1.4
	CERC8534	(6.0–)7.0–9.0(–10.0) × (2.0–)3.0–3.5(–4.0)	7.9 × 3.2	2.5	(3.0–)3.5–4.5(–5.0) × 2.0–3.0	4.0 × 2.5	1.6
	CERC8539	(5.5–)6.5–10.5(–12.5) × (2.5–)3.0–4.0(–4.5)	8.4 × 3.4	2.5	3.0–4.0(–4.5) × 2.0–3.0	3.6 × 2.5	1.4
	Average^d	(5.5–)6.5–9.5(–12.5) × (2.0–)3.0–4.0(–4.5)	7.9 × 3.3	2.4	(2.5–)3.0–4.0(–5.0) × 2.0–2.5(–3.0)	3.7 × 2.4	1.5

Tab.5 Primary conidial and secondary conidia measurements of three Quambalaria species identified in this study

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[1]	Yaojian XIE, Roger J. ARNOLD, Zhihua WU, Shuaifei CHEN, Apeng DU, Jianzhong LUO. Advances in eucalypt research in China[J]. Front. Agr. Sci. Eng. , 2017, 4(4): 380-390.

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