Phylogeny and taxonomy of Cinnamomum (Lauraceae)

Abstract

Taxonomy of Cinnamomum Schaeff. of Lauraceae remains problematic because recent phylogenetic studies have suggested that this genus is not monophyletic. In this study, we assembled three sequence matrices including plastomes (datamatrix I), nrITS sequences alone (datamatrix II), and nrITS plus plastid psbA‐trnH sequences (datamatrix III) of the Cinnamomum‐Ocotea complex of Lauraceae and conducted a new phylogenetic study with thusfar the most extensive species sampling of the Cinnamomum‐Ocotea group. We determined that the Old World Cinnamomum is diphyletic: sect. Camphora Meisn. is sister to Sassafras J.Presl and sect. Cinnamomum is sister to the African Kuloa Trofimov & Rohwer. A recent study indicated that characters of leaf micromorphological anatomy can define the two clades: one possessing reticulate periclinal and the other having non‐reticulate periclinal walls. As result, we divided the genus Cinnamomum of Lauraceae into two genera, i.e., Cinnamomum and Camphora Fabr. The generic name Cinnamomum is retained for those species mainly having reticulate periclinal epidermal cell walls, inconspicuous non‐perulate terminal buds and usually tripliveined leaves; the oldest generic name, Camphora, is applied to the second group which contains those species mainly possessing non‐reticulate periclinal epidermal cell walls, prominent perulate terminal buds and pinnately‐veined leaves. A census of the species and their type specimens listed under Cinnamomum in Asia resulted in the transfer of 18 species to Camphora, including 15 new combinations.

INTRODUCTION

The Lauraceae constitute the largest family of the order Laurales which is sister to the Magnoliales (The Angiosperm Phylogeny Group, 2016). This family contains more than 3000 species in ca. 55 genera that are mostly woody and widely distributed in the pantropics with tropical America and Australasia as the diversity centers (Renner, 2011). Taxonomy of the Lauraceae has been notorious due to imperfectly known species with either flower or fruit characters unknown, overlapping variation and parallel evolution of morphological characters, and poorly represented specimens in herbaria. Taxonomic delimitation of many generic groups has been controversial (van der Werff, 2001), e.g. Beilschmiedia group (Li et al., 2020; Yang et al., 2012), Persea group (Li et al., 2011; Mo et al., 2017), Ocotea group (Penagos Zuluaga et al., 2021; Trofimov et al., 2019, 2022), and Cinnamomum group (Gang et al., 2021; Huang et al., 2016; Rohde et al., 2017). The genus Cinnamomum Schaeff. is probably the most difficult one of the family Lauraceae because early researchers usually studied materials including detached leaves of uncertain origin that were mostly picked from immature plants (viz. cinnamon, Kostermans, 1970, 1985).

Cinnamomum is defined by a set of morphological characters including evergreen trees or shrubs, opposite and triplinerved or alternate and pinninerved leaves, paniculate-cymose inflorescences, bisexual and trimerous flowers, nine fertile stamens in three whorls with the two outer whorls introrse and the innermost whorl of stamens extrorse with normally 4-locular anthers, staminodes of the fourth whorl well developed, and fruits with a cupule (Lorea-Hernández, 1996; Rohwer, 1993; van der Werff, 2001). These morphological characters imply controversial systematic positions of Cinnamomum (Bentham & Hooker, 1880; Kostermans, 1957, 1986; Meissner, 1864; Pax, 1891; Rohwer, 1993; Van der Werff & Richter, 1996). Kostermans (1957) classified the genus Cinnamomum together with Ocotea Aubl., Actinodaphne Nees, Sassafras J. Presl, Umbellularia Nutt. and Dicypellium Nees & Mart. in the subtrib. Cinnamomineae of the trib. Cinnamomeae. Rohwer (1993) classified the family Lauraceae into informal groups, and ascribed the genus Cinnamomum to the Ocotea subgroup of the Ocotea group together with Neocinnamomum H.Liou, Aiouea Aubl., Endlicheria Nees, Rhodostemonodaphne Rohwer & Kubitzki, Ocotea, Nectandra Rol. ex Rottb., Pleurothyrium Nees. Van der Werff and Richter (1996), however, included Ocotea, Nectandra, Aniba Aubl., Licaria Aubl., Pleurothyrium Nees, Cinnamomum, Persea Mill., Phoebe Nees, and Dehaasia Blume in the trib. Perseeae according to types of inflorescences.

Phylogenies based on nrITS, 26S, trnL-trnF, psbA-trnH, trnT-trnL, and rpl16 sequences have suggested that the genus Cinnamomum belongs to the Cinnamomeae, but relationships of the Cinnamomeae were badly resolved with regard to Laureae (Chanderbali et al., 2001; Rohde et al., 2017). Based on a plastome phylogeny, Song et al. (2020) indicated a division of the family Lauraceae into six tribes with the genus Cinnamomum belonging to the trib. Laureae; that study did not sample the Mezilaurus group and the trib. Laureae is a mixture including the Persea group, the Cinnamomum group, and the Litsea group. Liu et al. (2021) suggested that the family Lauraceae should be classified into eight tribes according to recent phylogenetic studies, viz. Hypodaphnideae, Cryptocaryeae, Cassytheae, Neocinnamomeae, Caryodaphnopsideae, Perseeae, Cinnamomeae and Laureae (no samples of the Mezilaurus group), with the genus Cinnamomum included in the trib. Cinnamomeae.

The genus Cinnamomum was formerly considered to contain 350 species that are amphi-Pacific (Lorea-Hernández, 1996; Rohwer, 1993; van der Werff, 2001). Recent phylogenetic and taxonomic studies have transferred the American species to Aiouea (Rohde et al., 2017), so the remaining Old World Cinnnamomum now contains 247 species (). Species of Cinnamomum are usually subdivided into two groups: one group including species having alternate and pinnately veined leaves, domatia present in the axils of lateral veins and middle veins, and perulate terminal buds (Figure 1); and the other group containing species possessing opposite/subopposite and tripliveined leaves lacking domatia in the axils of lateral veins, and non-perulate terminal buds (Figure 2; Lorea-Hernández, 1996; Huang et al., 2016). The two groups are normally ranked as two sections, i.e. sect. Camphora Meisn. (syn.: sect. Malabathrum Meisn.) and Cinnamomum (Hooker, 1886; Kostermans, 1986; Li et al., 1982; Lorea-Hernández, 1996; Meissner, 1864) Nees (1831, 1836), however, separated the two groups into two genera: Camphora and Cinnamomum. Besides the macromorphological differences, the two groups are also different in characters of the upper leaf epidermis: (1) the cell shape is regular in sect. Camphora, but irregular in sect. Cinnamomum; (2) the anticlinal wall is straight or nearly so in sect. Camphora, but sinuous in sect. Cinnamomum; and (3) the periclinal wall is smooth in sect. Camphora, but reticulate in sect. Cinnamomum (Gang et al., 2021).

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Relationships of the Cinnamomum-Ocotea complex have not been resolved. Trofimov and Huang et al. (2016) suggested that sect. Cinnamomum is sister to the Neotropical clade, whereas Rohde et al. (2017) indicated that sect. Camphora is sister to the Neotropical clade. Trofimov and Rohwer (2020) suggested that the Old World Cinnamomum is diphyletic or paraphyletic as well, with sect. Cinnamomum sister to Kuloa Trofimov & Rohwer and sect. Camphora sister to Sassafras J. Presl. Other recent studies considered the genus Cinnamomum as paraphyletic with regard to Sassafras (Liu et al., 2021; Song et al., 2020; Trofimov et al., 2022), which may be attributable to sampling bias, none of them included Kuloa. The phylogeny of Cinnamomum is thus not well resolved, so further phylogenetic studies are necessary to determine the monophyly of Cinnamomum, and the genus should be further subdivided if confirmed to be polyphyletic. As a result, our target here is to (1) reconstruct a resolved phylogeny of the Cinnamomum–Ocotea complex with a broad sampling of the genus Cinnamomum using separate and concatenated sequence matrices including plastomes, nrITS and psbA–trnH sequences, and (2) conduct a new synoptic taxonomy of sect. Camphora if the polyphyly of the genus is confirmed.

MATERIALS AND METHODS

A new phylogeny of the Cinnamomum group was conducted using all available complete chloroplast genome/plastomes (CPG) and two commonly used markers including nrITS and psbA–trnH in the family Lauraceae. To resolve the phylogeny of Cinnamomum, we also included species samples of the Cinnamomum-Ocotea complex. Alseodaphne semecarpifolia Nees, Machilus thunbergii Siebold & Zucc., Persea americana Mill., and Phoebe sheareri Gamble were chosen as outgroups. Sequences were obtained from GenBank () and LCGDB () (Appendix A, last search 22 March 2022), aligned in MAFFTT (Katoh et al., 2017) using Auto and Localpair model for CPG and the two markers respectively, then adjusted and edited manually in BioEdit (Hall, 1999). Ambiguously aligned fragments of CPG were removed with Gblocks using default setting (Talavera & Castresana, 2007) and gap sites of nrITS and psbA-trnH sequences were deleted with trimAl using “-automated1” (Capella et al., 2009). Totally, we assembled three matrices: complete plastome sequences (datamatrix I), nrITS (datamatrix II), and a datamatrix including nrITS and psbA-trnH (datamatrix III). The two markers of datamatrix III were concatenated using PhyloSuite (Zhang et al., 2020). A best-fit or partition model of all matrices was computed with ModelFinder (Kalyaanamoorthy et al., 2017). For phylogenetic studies, maximum likelihood (ML) analyses were conducted in IQ-TREE (Nguyen et al., 2014), bootstrap values were obtained using Ultrafast Bootstrap for 5000 and 1000 times for the datamatrix I and other two datamatrices separately (Minh et al., 2013); Bayesian inferences (BI) were conducted in MrBayes (Ronquist et al., 2012) with the following parameters: generations: 12,000,000, sampling frequency: 6000, and burnin: 25.0%. Phylogenetic trees were viewed and edited in ITOL (Letunic & Bork, 2021), and improved in Adobe Illustrator 2020.

RESULTS

We finally obtained 38 plastomes, 324 nrITS and 238 psbA-trnH sequences. Statistic data of the three datamatrices and their best-fit models were listed in Table 1. Our plastome phylogeny (datamatrix I) contained four ingroup clades due to the lack of plastome sequences of Kuloa (Figure 3a). Our phylogenetic trees based on nrITS alone (datamatrix II) or nrITS plus psbA-trnH (datamatrix III) resulted in five large clades (Figures 3b,c and 4): Clade I including the American genera of the Cinnamomum-Ocotea complex; Clade II comprising sect. Camphora s.s. (excluding C. chago B.S.Sun & H.L.Zhao, C. longipetiolatum H.W. Li, and C. saxatile H.W. Li, here defined); Clade III containing the deciduous genus Sassafras; Clade IV including sect. Cinnamomum s.l. (including C. chago, C. longipetiolatum, and C. saxatile of sect. Camphora s.l.); and Clade V encompassing the African Kuloa. These phylogenetic trees based on different datamtrices gave rise to similar relationships of the five large clades, the genus Cinnamomum was polyphyletic, sect. Camphora s.s. was sister to Sassafras, and sect. Cinnamomum s.l. was sister to Kuloa. Relationships within the two sections of Cinnamomum were not resolved. To show morphological differences of the two groups of Cinnamomum, we mapped both macro- and micro-morphological characters on the combined tree based on nsITS plus psbA-trnH sequences (Figure 4).

TABLE 1 Statistics of sequence information for phylogeny of this study

Item	nrITS	nrITS + psbA-trnH	psbA-trnH	Plastome
Aligned length (nt)	867	1511	644	152,510
Variable sites (nt)	458	743	285	3611
Parsimony informative sites (nt)	310	486	176	1690
V%	52.83%	49.17%	44.25%	2.37%
P%	35.76%	32.16%	27.33%	1.11%
Model (ML)	TIM + F + I + G4	Partitioned	–	K3Pu + F + I + G4
Model (BI)	GTR + F + I + G4	Partitioned	–	GTR + F + I + G4

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DISCUSSION

Our new phylogenies using nrITS alone and nrITS plus psbA-trnH sequences included thusfar the most extensive species sampling of the Cinnamomum-Ocotea complex. We have identified five large clades including the Ocotea group (Clade I), sect. Camphora (Clade II), Sassafras (Clade III), sect. Cinnamomum (Clade IV), and Kuloa (Clade V). The phylogenetic position of the Ocotea group is different between our new phylogeny and a few previous studies (Huang et al., 2016; Rohde et al., 2017; Trofimov & Rohwer, 2020): the Ocotea group is sister to a clade including three subclades (sect. Camphora, sect. Cinnamomum and Sassafras) or to two subclades (sect. Camphora and Sassafras) in our new phylogeny but sister to a clade containing sect. Cinnamomum plus Kuloa (the African Ocotea) in Huang et al. (2016), and sister to sect. Camphora plus Sassafras in Trofimov and Rohwer (2020). Rohde et al. (2017) suggested additional relationships, i.e. Sassafras alone is sister to a large clade including the Ocotea group plus Cinnamomum, in the large clade the Ocotea group and sect. Camphora forms a subclade sister to sect. Cinnamomum; their phylogenetic trees possess very low bootstrap values. However, our plastome phylogenetic tree shows similar topology to that of Trofimov et al. (2022) that the Ocotea group is sister to a clade including sect. Cinnamomum and sect. Camphora plus Sassafras. Our new phylogenetic results clearly suggest that the Old World Cinnamomum species are diphyletic and represent two separate groups, which is consistent with recent studies using representative species sampling (Huang et al., 2016; Rohde et al., 2017; Trofimov & Rohwer, 2020). Our plastome phylogeny indicates that the genus Cinnamomum is paraphyletic with respect to Sassafras, which agrees with the result of Trofimov et al. (2022); this relationship is probably caused by the lack of sampling in the African Kuloa and the incongruence between cpDNA and nuclear DNA phylogenies; no plastomes of Kuloa are available at present. Taken together, we conclude that the genus Cinnamomum is diphyletic.

For a new classification, we also considered macromorphology and micromorphology. Macromorphological characters are largely consistent with the phylogenetic results, e.g. buds perulate or not, leaves alternate or opposite, pinnately veined or tripliveined, domatia presence in axil of lateral veins (Figure 5), except for C. chago, C. longipetiolatum, and C. saxatile. A recent study of leaf epidermal micromorphology in the Old World Cinnamomum species by Gang et al. (2021) found two types of micromorphology that were clade-specific and highly predictive. The periclinal wall ornamentation coincides perfectly with the phylogenetic results seen here, i.e., sect. Camphora s.s. possessing a non-reticulate periclinal wall and sect. Cinnamomum s.l. having a reticulate periclinal wall (Gang et al., 2021; and our study here). Considering the congruence of macromorphological, micromorphological and phylogenetic results, Cinnamomum as currently circumscribed is therefore best divided into two genera.

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Cinnamomum sect. Cinnamomum s.l. embraces the generic type: C. verum J. Presl (syn.: C. zeylanicum Blume), and thus, should retain the generic name and a different generic name be given to sect. Camphora s.s. under Art. 10.8 of the Shenzhen Code (Turland et al., 2018). There are several generic names listed under synonymy for Cinnamomum s.l. (e.g., Li et al., 1982, 2008; Rohwer, 1993), including Camphora Fabr., Cecidodaphne Nees, Parthenoxylon Blume and Temmodaphne Kosterm. Cecidodaphne Nees is based on C. glaucescens Nees (≡Cinnamomum glaucescens [Nees] Hand.-Mazz.) and should be considered as a synonym of Cinnamomum in the narrow sense, because the type species bears tripliveined leaves and belongs to sect. Cinnamomum. Similarly, the type specimen of Temmodaphne (T. thailandica Kosterm.) also has triplinerved, sub-opposite leaves (Kostermans, 1973) suggesting that it also belongs in Cinnamomum s.str. Parthenoxylon is based upon P. porrectum (Roxb.) Blume, and treated as a synonym of C. parthenoxylon (Jack) Meisn. by the Flora of China (Li et al., 2008), so is potentially available for the non-Cinnamomum clade. However, Camphora (Fabricius, 1759) has priority over Parthenoxylon (Blume, 1849–1851). As a result, we transfer those species usually with alternate and pinnately veined leaves, domatia present in axil of lateral veins, perulate buds, and non-reticulate periclinal walls to Camphora.

TAXONOMIC TREATMENT

Cinnamomum

Schaeff., Bot. Exped. 74. Oct-Dec 1760 (nom. cons.). Type: C. verum J. S. Presl (in Berchtold & J. S. Presl, Prir. Rostlin 2: 36. 1825) (Laurus cinnamomum L.)

= Cecidodaphne Nees, Wall. Pl. Asiat. Rar. 3: 72. 1831. Type: C. glaucescens C. G. D. Nees.

Diagnosis

Buds usually not perulate. Leaves usually subopposite and tripliveined, rarely alternate and pinnately veined, domatia absent, adaxial epidermal cells irregular in shape, anticlinal walls sinuous, rarely straight, periclinal walls reticulate. Inflorescences paniculate with cymes bearing strictly opposite lateral flowers. Flowers bisexual with nine fertile stamens, plus three staminodes with conspicuous cordate or sagittate heads in the fourth androecial whorl. Fruits cupulate with tepals at least partially persistent. Pedicels turbinate.

Distribution

Tropical to subtropical Asia.

Remarks

Several species were placed previously into sect. Camphora because they have seemingly pinnately veined leaves, e.g., Cinnamomum chago, C. longipetiolatum, and C. saxatile, but our molecular study suggests that these species belong to Cinnamomum (Figures 3 and 4). Similarly, Gang et al. (2021) also suggested that C. saxatile belongs to the former sect. Cinnamomum as it possesses reticulate periclinal walls. Sun and Zhao (1991) noted that leaves of C. chago are pinnately veined with 7–9 pairs of lateral veins, the proximal pair starting from the base of leaf blade and appearing subtriveined, suggesting that the leaf venation of the species is probably triveined. However, leaf micromorphology should be examined for these three species (and other Cinnamomum-like taxa, such as Temmodaphne thailandica) before their taxonomic position can be confirmed. Therefore, we retain these species in Cinnamomum for now, pending further study.

Camphora

Fabr., Enum. 218. 1759. Type: C. officinarum Nees in Wallich, Pl. Asiat. Rar. 2: 72. 1831 ≡ Laurus camphora L., Sp. Pl. 1: 369. 1753

= Parthenoxylon Blume, Mus. Bot. 1: 322. 1851. Lectotype: Laurus parthenoxylon W. Jack (vide Pfeiffer, Nom. 2: 598. 3 Oct 1873).

Diagnosis

Buds usually perulate. Leaves alternate and pinnately veined or weakly tripliveined, domatia usually present in axils of lateral veins, adaxial epidermal cells polygonal, anticlinal walls straight or nearly so, periclinal walls smooth and non-reticulate. Inflorescences paniculate with cymes bearing strictly opposite lateral flowers. Flowers bisexual with nine fertile stamens, plus three staminodes with conspicuous cordate or sagittate heads in the fourth androecial whorl. Fruits cupulate with tepals not or partially persistent. Pedicels turbinate.

Distribution

Tropical to subtropical Asia, but mostly distributed in the Northern Hemisphere (Soh, 2011). In China, there are ca. 18 species of Camphora; Kochummen (1989), Soh (2011) and de Kok (2019) recorded only one species of Camphora (viz. Cinnamomum porrectum (Blume) Kosterm., synonym of Camphora parthenoxylon (Jack) Nees in this treatment) in the Tree Flora of Malaya, in Borneo and in Peninsular Malaysia and Singapore respectively; there is no species with pinnately veined leaves in southern India (Kostermans, 1985).

Camphora bodinieri

(H. Lév.) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Cinnamomum bodinieri H. Lév., Repert. Spec. Nov. Regni veg. 10: 369. 1912 — Type: CHINA. Guizhou (贵州, ‘Kouy-Tchéou’): Near Guiyang (贵阳, ‘Kouy-Yang’), “bois de la pagode de Lan-Yo-Chan”, 15 Jun 1899, Bodinier 2622 (holotype: E00386445!; isotypes: P01978724 [photo!], P01978725 [photo!]; fragm. K000778561!, fragm. A00041222 [photo!], with photo of holotype).

= Cinnamomum glanduliferum var. longipaniculatum Lec., Nouv. Arch. Mus. Hist. Nat., sér. 5, 5: 74. 1913 — Type: CHINA. Chongqing (重庆): Chengkou (城口, as ‘district de Tchen-kéou-tin’). ‘Moùng Moùng Ky’, alt. 1400 m, Farges 894 (lectotype: P01978761!, designated here; isolectotypes: K000778562!, P01964399!, P01964400!, P01964401!, P01978730!, P01978764!).

= Cinnamomum inunctum var. fulvipilosum Y.C. Yang, J. West China bord. Res. Soc. 15 (Ser. B): 73. 1945 — Type: CHINA. Guizhou (贵州, as ‘Kweichow’), Zunyi (遵义), ‘Liang-Feng-Yah’, rocky slope near farmhouse, alt. 1000 m, 1 Aug 1931, A. N. Steward, C. Y. Chiao (焦启源), and H. C. Cheo 143 (holotype: PE00028456 [photo!]; isotypes: K000778560, N102060173 [photo!], NAS00188664 [photo!], P00757069, PE00189210 [photo!], PE00028450 [photo!], PE00028457 [photo!]).

Distribution: Guizhou, Hubei, Hunan, Shaanxi, Sichuan, Yunnan.

Camphora brachythyrsa

(J. Li) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Cinnamomum brachythyrsum J. Li, Acta Bot. Yunnan 18: 53. 1996 — Type: CHINA. Yunnan (云南): Wenshan (文山), Laojunshan (老君山), May 1993, Y.M. Shui 003072 (holotype: KUN).

Distribution: Yunnan.

Camphora chartophylla

(H.W. Li) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Cinnamomum chartophyllum H.W. Li, acta Phytotax. Sin. 13 (4): 491975 — Type: CHINA. Yunnan (云南), Menglun (勐仑), Sheng-Ji Pei (裴盛基) 59–10384 (holotype: KUN; Isotype: fragm. L0035751)

Distribution: Yunnan.

Camphora foveolata

(Merr.) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Beilschmiedia foveolata Merr., J. Arnold arbor. 19(1): 30. 1938 ≡ Cinnamomum foveolatum (Merr.) H.W.Li & J.Li, Fl. China 9: 170. 2008 ≡ Litsea foveolata (Merr.) Kosterm., Reinwardtia 8: 98. 1970, not yen C. Yang & P. H. Huang (1978) — Type: VIETNAM. Tonkin, Chapa, alt. 1700 m, Aug., 1930, Petelot 5580 (A, BO, NY, P)

Machilus camphorata [“camphoratus”] H. Lév., Repert. Spec. Nov. Regni Veg. 9: 460. 1911 ≡ Alseodaphne camphorata (H.Lév.) C.K.Allen, J. Arnold Arbor. 17(4): 326 (1936) ≡ Alseodaphne caudata Lec. Nouv. Arch. Mus. Hist. Nat., sér. 5, 5: 97–98. 1913 ≡ Cinnamomum caudifer Kosterm., Reinwardtia 8: 35. 1970 — Type: CHINA. Guizhou (贵州, as ‘Kouy-Tcheou’): Guiding (贵定), Pingfa, (平伐, as ‘Pinfa’), Apr 1908 [on K000778566] or 7 May 1903 [on K000778567], Cavalerie 1002 (holotype: E00386438; isotypes: K000778566, K000778567, L0035749, P00757048 [holotype of Alseodaphne caudata], fragm. A00415028).

Distribution: Guizhou, Yunnan; VIETNAM.

Camphora glandulifera

(Wall.) Nees, Pl. Asiat. Rar. 2: 72. 1831 ≡ Laurus glandulifera Wall., Trans. Med. Soc. Calcutta 1: 45, 51, pl. 1. 1825 ≡ Cinnamomum glanduliferum (Wall.) Meisn. In A. P. de Candolle, Prodr. 15 (1): 25 (1864) ≡ Camphora rougierii var. glandulifera (Nees) Lukman., Nomencl. Icon. Cannel. 23. 1889, nom. Illeg. — Type: NEPAL. Mount Shivapuri [“ad Sheopore mont.”], 1821, Wallich 2601 (holotype: K001116542; Isotypes: B100277066, B100277067, BM000880671, GZU000253941, LE00012754, Fragm. A00041270, With photo of K001116542)

= Machilus mekongensis Diels, Notes Roy. Bot. Gard. Edinburgh 5 (25): 244: 1912 — Type: CHINA. Yunnan (云南): Weixi (维西), Cikai Town (茨开), Dong-Shan (东山), ‘Shupa valley. Shrubby hillsides. Alt. 11,000 ft. Yangtse-Mekong divide, Tibet’, 1904, Forrest 370 (lectotype: E00386432, designated here).

= Cinnamomum cavaleriei H. Lév., Repert. Spec. Nov. Regni Veg. 10: 370. 1912 — Type: CHINA. Guizhou (贵州, as ‘Kouy-Tcheou’): Pingfa, (平伐, as ‘Pin-Fa’), 23 Jun 1903, Cavalerie 1084 (holotype: E00386433; isotypes: E00386434, K000778586, fragm. A00041223).

= Machilus dominii H. Lév., Repert. Spec. Nov. Regni Veg. 13: 174. 1914 ≡ Cinnamomum dominii (H. Lév.) C. F. Ji, J. Nanjing For. Univ. 25(3): 76. 2001 — Type: CHINA. Yunnan (云南): Kunming (昆明), ‘Forêts de Ku-Long-Tchang’ (古龙场), alt. 800 m, Jul 1912, Maire 35 (holotype: E00386435; isotypes: BM000950907, fragm. A00041227).

Distribution: Guizhou, Sichuan, Yunnan, Xizang; BHUTAN, INDIA, MALAYSIA, MYANMAR, NEPAL.

Remarks: The specimen NY00355188 labeled as isotype of Laurus glandulifera is not an isotype. It has been collected by E. Meyer in Java in 1842, and the text of Wallich was added in quotes by Meisner.

We were unable to locate the second syntype of Machilus mekongensis: CHINA. Yunnan (云南): Weixi (维西), Cikai Town (茨开), Dong-Shan (东山), ‘Mekong-Salween divide behind Tsekou mission, Tibet’, 1904, Forrest 372.

Camphora illicioides

(A. Chev.) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Cinnamomum illicioides [“ilicioides”] A. Chev., bull. Écon. Indochinen. s. 20: 855. 1918 — Type: VIETNAM. “Phu-tho, Vinh-yên, montagnes du Tam-dao, etc.”, Phu-tho: Trung Giáp forest reserve, 29–30 May 1918, Fleury 37,993 (holotype: P00757044; Isotypes: K000350907, L0035795)

Distribution: Guangxi, Hainan; THAILAND, VIETNAM.

Camphora longepaniculata

(Gamble) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Cinnamomum inunctum var. longepaniculatum Gamble in C. S. Sargent, Pl. Wilson. 2: 69. 1916 ≡ Cinnamomum longepaniculatum (Gamble) N. Chao ex H.W. Li, Acta Phytotax. Sin. 13 (4): 48, f. 2. 1975 — Type: CHINA. Sichuan (四川): Ya'an (雅安, as ‘Yachou Fu’), 600–1000 m, Jun 1980, Wilson 3710 (holotype: A00041232; Isotypes: BM000950908, HBG509751, US00099081)

Distribution: Sichuan.

Camphora micrantha

(Hayata) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Machilus micrantha Hayata, icon. Pl. Formosan. 2: 130. 1912 ≡ Cinnamomum micranthum (Hayata) Hayata, Icon. Pl. Formosan. 3: 160. 1913 — Type: CHINA. Taiwan (台湾): Xinbei City (新北市), Sanxia (插角[三峡], 大豹), as “Sankakuyu, Taihyo”, Kanehira s.n., Jun 1912 (holotype: TI No. 02537)

= Cinnamomum kanehirae [“kanehirai”] Hayata, Icon. Pl. Formosan. 3: 159. 1913 ≡ C. micranthum f. kanehirai (Hayata) S. S. Ying, Mém. Coll. Agric. Natl. Taiwan Univ. 25 (1): 108. 1986 — Type: CHINA. Taiwan (台湾): Miaoli (苗栗), Nanzhuang (南庄, 加里前山), as “Nanshoshicho, Kalizenzan”, alt. 4000 ft., Oct. 1912, Kanehira s.n. (holotype: TI2456).

= Cinnamomum xanthophyllum H. W. Li, Acta Phytotax. Sin. 13(4): 47. 1975 — Type: CHINA. Guangdong (广东): Xinfeng (新丰), alt. 650 m, L. Deng (邓良) 8043 (holotype: KUN; isotypes: IBK00004390, IBSC0046454, SZ00162655, AU034001, PE00189952).

Distribution: Fujian, Guangdong, Guangxi, Guizhou, Hainan, Jiangxi, Taiwan; VIETNAM.

Camphora migao

(H.W. Li) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Cinnamomum migao H.W. Li, Acta Phytotax. Sin. 16 (2): 90, pl. 7, f. 1. 1978 — Type: CHINA. Yunnan (云南): Funing (富宁), alt. 500 m, H. T. Tsai (蔡希陶) 58–9048 (holotype: KUN; Isotypes: IBK00200070, LBG00072386)

Distribution: Guangxi, Yunnan.

Camphora mollifolia

(H.W. Li) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Cinnamomum mollifolium H.W. Li, Acta Phytotax. Sin. 13 (4): 45, f. 1. 1975 — Type: CHINA. Yunnan (云南): Menghai (勐海), Y. H. Li (李延辉) 60–11,664 (holotype: KUN48456; Isotype: Fragm. L0035915)

Distribution: Yunnan.

Camphora officinarum

Nees, Pl. Asiat. Rar. 2: 721831 ≡ Laurus camphora L., Sp. Pl. 1: 369. 1753 ≡ Persea camphora (L.) Spreng., Syst. Veg.2: 268. 1825 ≡ Cinnamomum camphora (L.) J. Presl, Prir. Rostlin 2: 36, pl. 8. 1825 ≡ Camphora officinalis Steud., Nomencl. Bot., ed. 2 (Steudel) 1: 271. 1840 ≡ Camphora camphora (L.) H. karst., Deut. Fl. 504. 1881, nom. inval. — Type: Japan. Locality and date not indicated, collector not indicated (lectotype: LINN 518.7 [photo!], designated by Kostermans, 1978)

= Camphora japonica Garsault, Descr. Vert. Usag. 719. Pl. 1: 61, t. 84. 1767 — Holotype: Japan. no collection indicated; plate 84 may be accepted as type.

= Camphora officinarum var. glaucescens A. Braun, Verh. Vereins Beförd. Gartenbaues Königl. Preuss. Staaten 21: 77 (1853); Cinnamomum camphora var. glaucescens (A. Braun) Meisn. in A. P. de Candolle, Prodr. 15 (1): 24. 1864 — Type: cultivated, presumably in Berlin (holotype: B, possibly destroyed; isotype: P01991848).

= Cinnamomum camphora fo. parvifolia Miq., Ann. Mus. Bot. Lugduno-Batavi 2: 195. 1866 — Type: Japan. Nagasaki: 1862–1863, Oldham 704 (holotype: L0308679 fruiting with Miquel's handwriting; isotype P00757059, probable type of Camphora humboldtii, see below).

= Camphora oldhamii Lukman., Nomencl. Icon. Cannel. 23. 1889 — probable type (fide Kostermans): CHINA. Taiwan (台湾): 1864, Oldham 44? [interpreted as 4425 in P; could equally well be 447 or 449] (P00757057).

= Cinnamomum camphora var. nominale Hayata, J. Coll. Sci. Imp. Univ. Tokyo 22: 349. 1906 ≡ C. nominale (Hayata) Hayata, Icon. Pl. Formosan. 3: 160. 1913 — Holotype: CHINA. Taiwan (台湾): Hengchun [“Koshun”], 1905, Kawakami (not found); neotype (designated here): CHINA. Taiwan (台湾): Pingdong (屏东), Ken-Ding-Park (垦丁公园), Kuraru (龟子角, as ‘Koshun’), May 30th, 1912, B. Hayata s.n. (TI no. 02459 [photo!]; isoneotypes: TI nos. 02460 [photo!], 02461 [photo!], 02462 [photo!]).

= Cinnamomum taquetii H. Lévl., Feddes Repert. 10: 370. 1912 — Type: SOUTH KOREA. Jeju Island (济州岛, as ‘Quelpaert’), Daejeong-eup (大静, ‘in silvis Taitpjeng’), Jul 1909, Taquet 3159 (Lectotype: E00386436 [photo!], designa; isolectotypes: KYO, TI).

= Cinnamomum camphoroides Hayata, Icon. Pl. Formosan. 3: 158. 1913 — Type: CHINA. Taiwan (台湾): Pingdong (屏东), Koshun (恒春), N.Konishi s.n. (holotype: TI 2048; isotype: L0035885).

= Cinnamomum nominale var. lanata Nakai, Fl. Sylv. Kor. 22: 301939 — Type: CHINA. Taiwan (台湾): Hualiangang (花莲港, as ‘Kwarenko’), date and collector not indicated (holotype: TI no. 02463).

= Cinnamomum simondii Lecomte, J. Arnold Arbor. 20: 45 (1939). — Type: CHINA. Guangxi (广西): Longzhou (龙州, as ‘Long Tchéou’), without date, Simond 190 (holotype: P00757021; isotypes: K000778667, L0035964; photo of holotype, A00041278).

= Cinnamomum camphora var. cyclophyllum Nakai, J. Jap. Bot. 19: 369. 1943 — Type: SOUTH KOREA. Jeju Island (济州岛, as ‘Saisyuto’), ‘in sylvis montis Kanrasan’ (汉拏山), 10 Nov. 1917, Takahasi s.n. (holotype: TI no. 02444).

= Cinnamomum camphora var. linaloolifera Y. Fujita, Bot. Mag. (Tokyo) 65: 2451952 ≡ Cinnamomum camphora f. linaloolifera (Y. Fujita) Sugim., New Keys Jap. Trees 459. 1961 — Type: CHINA. Taiwan (台湾): ‘Hab. Formosa’, collector and date not indicated (not seen).

Distribution: wide spread in southern China; Japan, Korea, Vietnam.

Camphora parthenoxylon

(Jack) Nees in wall. Pl. Asiat. Rar. 2: 72. 1831 ≡ Laurus parthenoxylon Jack, Malayan Misc. 1(5): 28. 1820 ≡ Camphora parthenoxylon (Jack) Nees in Wall., Pl. Asiat. Rar. 2: 72. 1831 ≡ sassafras parthenoxylon (Jack) Nees, syst. Laur. 491. 1836 ≡ Cinnamomum parthenoxylon (Jack) Meisn. In A. Candolle Prodr. 15(1): 26. 1864 — Type: [INDONESIA]. Sumatra, ‘kayo Gadis’, herb. Roxburgh s.n. (holotype: BR0000005931088; Isotype: BR0000005931132)

≡ Laurus porrecta Roxb., Hort. Beng. 30. 1814, nom. Inval. ≡ Camphora porrecta (Roxb.) Voigt, Hort. Suburb. Calc. 308. 1845, nom. inval. ≡ Parthenoxylon porrectum Blume, Mus. Bot. 1: 323. 1851 ≡ Cinnamomum porrectum (Blume) Kosterm. J. Sci. Res. (Jakarta) 1(5): 126. 1952 — Type: [INDONESIA. Sumatra or India, cult. Hort. Bot. Calcutta] Roxburgh s.n. (neotype: BR0000005931088, designated by Kostermans, 1970, second-step by Turner, 2013; isoneotype: BR0000005931132).

= Cinnamomum barbatoaxillatum N. Chao in Fl. Sichuan. 1: 36, 459. 1981 — Type: CHINA. Sichuan (四川): Yibin (宜宾), fruit, N. Zhao 2918 (holotype: SCFI; isotypes: IBK00345678, KUN0101453).

Distribution: Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangxi, Sichuan, Yunnan; BHUTAN, CAMBODIA, INDIA, INDONESIA, LAOS, MALAYSIA, MYANMAR, NEPAL, PAKISTAN, THAILAND, VIETNAM.

Camphora philippinensis

(Merr.) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Machilus philippinensis Merr., Philipp. J. Sci. 1 (suppl. 1): 561906 ≡ Persea philippinensis (Merr.) Elmer, Leafl. Philipp. Bot. 2: 384. 1908 ≡ Cinnamomum philippinense (Merr.) C.E. Chang, Fl. Taiwan 2: 417. 1976 — Type: PHILIPPINES. Luzon: Prov. Bataan, Lamao River, mt. Mariveles, mar 1905, Meyer 2793 (lectotype: US00516627; Isolectotypes: NY00355328, NY00355329, NY00355330)

= Cinnamomum acuminatissimum Hayata, Icon. Pl. Formosan. 3: 157–158. 1913 ≡ Machilus acuminatissima (Hayata) Kaneh., Formosan Trees (rev. ed.) 219. 1936 ≡ Persea acuminatissima (Hayata) Kosterm., Reinwardtia 6 (2): 191. 1962 — Type: CHINA. Taiwan (台湾): Hualian (花莲), Taisho (大庄), March 26th, 1911, Furukawa s.n. (holotype: TI no. 02442; possible isotypes: K000778559, fragm. L0035683).

= Cinnamomum caudatifolium Hayata, Icon. Pl. Formosan. 5: 155, f. 54b. 1915 — Type: CHINA. Taiwan (台湾): Jiayi (嘉义县), Alishan [“Mt. Arisan”], between Karapin (Chaoliping交力坪) and Fenchihu [奋起湖, as “Funkiko”], near Shuisheliao [水车竂, as “Suisharyo”], [27] Mar. 1914, Hayata s.n. (holotype: TI no. 02450; isotype: fragm. L0035684).

Distribution: Taiwan; PHILIPPINES.

Remarks: We selected the collection Meyer 2793 (US00516627) as lectotype of Machilus philippinensis because it bears an original data sheet and apparently has been annotated by Merrill. The two other syntypes are: PHILIPPINES. Luzon: Prov. Bataan, Lamao River, Mt. Mariveles, Mar 1905, Whitford 1139 (K000778822, NY00355330, US00099164); PHILIPPINES. Luzon: Prov. Bataan, Lamao River, Mt. Mariveles, Apr 1905, Whitford 1220 (K000778823, NY00355328, US00516628). The collection Elmer 8184, labeled in several herbaria as type, has been cited by Elmer, but is not a type.

Camphora platyphylla

(Diels) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Machilus platyphylla Diels, Bot. Jahrb. Syst. 29: 348. 1900 ≡ Cinnamomum platyphyllum (Diels) Allen, J. Arnold arbor. 20: 46. 1939 — Lectotype (designated here): CHINA. Chongqing (重庆): Nanchuan (南川), native collectors, commun. Bock & von Rosthorn 1981 (A00041247; isolectotype: fragm. L0035914)

= Cinnamomum chengkouense N. Chao, Fl. Sichuan. 1: 459–460, f. 13. 1981 —Type: CHINA. Chongqing (重庆): Chengkou (城口), T. L. Dai (戴天伦) 102,187 (holotype: SZ; isotypes: PE00189181, IBK00190147).

Distribution: Chongqing, Sichuan.

Camphora purpurea

(H.G. Ye & F.G. Wang) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Cinnamomum purpureum H.G. Ye & F.G. Wang, Novon, 16: 439. 2006. — Holotype: CHINA. Guangdong (广东): Yangchun city (阳春市), Ehuangzhang mtn., safflower pond, ca. 300–800 m, 2 mar. 2002, ye Hua-gu & ye Yu-shi 6892 (IBSC; isotypes, IBSC)

Distribution: Guangdong.

Remarks: This name was treated as a synonym of Cinnamomum parthenoxylon (Jack) Meisn. in the Flora of China (Li et al., 2008). However, this species markedly differs from the latter in the purplish color of its branchlets, petioles, pedicel, and peduncles. We thus treat it as a separate species here.

Camphora rufotomentosa

(K.M. Lan) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Cinnamomum rufotomentosum K.M. Lan, Fl. Guizhou 2: 674. Pl. 32. 1984 — Holotype: CHINA. Guizhou (贵州): Xingyi (兴义), K.M. Lan 40 (GZAC).

Distribution: Guizhou.

Camphora septentrionalis

(Hand.-Mazz.) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Cinnamomum septentrionale Hand.-Mazz., Oesterr. Bot. Z. 85: 213–214. 1936 — Type: CHINA. Shaanxi (陕西): N side of tapa-Shan near Hanzhong (汉中, as ‘Hantschung’), Xiao-Nan-Hai, 800 m, May-Jun 1934, Fenzel 633 (holotype: W, probably destroyed in world war II; isotypes: fragm. A00246778, with photo of holotype [left image]; L0035916; PE00294191)

= Cinnamomum inunctum var. albosericeum Gamble in C. S. Sargent, Pl. Wilson. 2: 69. 1916 — Type: CHINA. Sichuan (四川): Mianzhu Xian [as “Mien-chu Hsien”, 绵竹县], 600 m, 19 May 1908, Wilson 3713 (holotype: A00041231; isotypes: B100277110, HBG509750, HUHA00041231 [photo!], IBSC0046892, IBSC0046891, L0035712, US00099080).

Distribution: Gansu, Shaanxi, Sichuan.

Camphora tenuipilis

(Kosterm.) Y. Yang, Bing Liu & Zhi Yang, comb. nov. ≡ Alseodaphne mollis W.W. Sm., notes Roy. Bot. Gard. Edinburgh 13: 153–154. 1921; ≡ Cinnamomum tenuipile (“tenuipilis”) Kosterm., Reinwardtia 8: 74. 1970 — Type : CHINA. Yunnan (云南): Shweli-Salween divide, in thickets. Lat. 25°30′N. Alt. 9000 ft., oct 1917, Forrest 16,021 (lectotype: E00123606, designated by Kostermans, 1970; isolectotype: K000350906).

Distribution: Yunnan.

Remarks: Alseodaphne mollis was based on two syntypes. The other syntype is: CHINA. Yunnan: Salween Valley, in open thickets. Lat. 25°6′ N. Alt. 4000 ft., Apr. 1917, Forrest 13,667 (E00123605; isosyntype: K000350905). When Kostermans (1970) transferred the species to Cinnamomum, he wrote “Typus: Forrest 16021 (E), syntypus: Forrest 13667 (E).” This may be interpreted as lectotypification in the sense of the fruiting specimen E00123606, even though E00123605 is the better flowering material.

AUTHOR CONTRIBUTIONS

Zhi Yang: Data curation (lead); formal analysis (lead); methodology (lead); software (lead); writing – original draft (equal). Bing Liu: Conceptualization (supporting); investigation (supporting); resources (lead); visualization (lead); writing – review and editing (equal). Yong Yang: Conceptualization (lead); funding acquisition (lead); investigation (lead); project administration (lead); supervision (lead); writing – original draft (lead); writing – review and editing (lead). David Kay Ferguson: Investigation (supporting); writing – review and editing (supporting).

ACKNOWLEDGMENTS

We are thankful to Jens G. Rohwer for his kind help on typification of the names, and constructive suggestions on an earlier version of this manuscript; we are also grateful to two anonymous reviewers for their valuable suggestions. This work was supported by the National Natural Science Foundation of China [31970205, 31770211, 31270238], National Natural Science Foundation of Jiangsu Province [BK20211279], and the Metasequoia funding of Nanjing Forestry University to YY.

CONFLICT OF INTEREST

The authors declare that there is no conflict of interest.

DATA AVAILABILITY STATEMENT

All data used in the study are included in this paper.

Appendix

APPENDIX A - Sequences obtained from GenBank for phylogenetic analysis

Taxon	nrITS	psbA-trnH	Plastome	Clade
Aiouea acarodomatifera	MF110006.1	–	–	I
Aiouea alainii	MF110007.1	–	–	I
Aiouea amoena	MF110008.1	MF137928.1	–	I
Aiouea chavarriana	MF110009.1	MF137929.1	–	I
Aiouea cinnamomoidea	AF272288.1	–	–	I
Aiouea dubia	MF110012.1	MF137932.1	–	I
Aiouea erythropus	AF272264.1	–	–	I
Aiouea formicaria	KX509823.2	KX509883.1	–	I
Aiouea grandifolia	MF110014.1	MF137934.1	–	I
Aiouea guianensis	AF272251.1	AF268780.1	–	I
Aiouea hammeliana	MF110016.1	MF137936.1	–	I
Aiouea haussknechtii	MF110019.1	MF137937.1	–	I
Aiouea hirsuta	KX509824.1	KX509884.1	–	I
Aiouea maya	MF110020.1	MF137940.1	–	I
Aiouea montana	MF110021.1	MF137941.1	–	I
Aiouea myristicoides	MF110022.1	MF137942.1	–	I
Aiouea obscura	MK507230.1	MK507298.1	–	I
Aiouea padiformis	KU139868.1	KU160284.1	–	I
Aiouea palaciosii	MF110023.1	MF137943.1	–	I
Aiouea pittieri	KU139836.1	–	–	I
Aiouea saligna	KX509821.1	KX509881.1	–	I
Aiouea sellowiana	MF110025.1	MF137945.1	–	I
Aiouea subsessilis	KU139889.1	KU160287.1	–	I
Aiouea tetragona	AF272265.1	AF268781.1	–	I
Aiouea tomentosa	MF110031.1	MF137951.1	–	I
Aiouea tomentosa	FM957804.1	–	–	I
Aiouea trinervis	MF110032.1	MF137952.1	–	I
Alseodaphne semecarpifolia	MG188583.1	–	NC_037491.1	Outgroup
Aniba affinis	MK507231.1	MK507299.1	–	I
Aniba canelilla	MW489499.1	–	–	I
Aniba excelsa	AF272255.1	–	–	I
Aniba firmula	MF110034.1	MF137954.1	–	I
Aniba heringeri	GQ480364.1	–	–	I
Aniba panurensis	AF272256.1	GQ428739.1	–	I
Aniba parviflora	MW489500.1	KX248005.1	–	I
Aniba rosaeodora	MW489501.1	MT679556.1	–	I
Aniba taubertiana	MK507233.1	KX248016.1	–	I
Aniba terminalis	MW489502.1	KX248008.1	–	I
Cinnamomum agasthyamalayanum	MH232437.1	–	–	IV
Cinnamomum appelianum	KP092853.1	KX546093.1	–	IV
Cinnamomum aromaticum		–	NC_046019.1	IV
Cinnamomum austrosinense	KU139818.1	KJ686727.1	–	IV
Cinnamomum austroyunnanense	KU139819.1	–	–	IV
Cinnamomum baileyanum	KU139820.1	KU160274.1	–	IV
Cinnamomum bejolghota	KX546412.1	EU153949.1	–	IV
Cinnamomum bodinieri	MH270478.1	MF137955.1	–	II
Cinnamomum burmanni	MF110036.1	MF137958.1	MW421305.1	IV
Cinnamomum camphora	KT248576.1	GU135428.2	MF421523.1	II
Cinnamomum celebicum	KU139829.1	–	–	IV
Cinnamomum chago	KU139830.1	KX546108.1	MN047449.1	IV
Cinnamomum chartophyllum	KU139832.1	–	MW421301.1	II
Cinnamomum chemungianum	MH232440.1	–	–	IV
Cinnamomum cordatum	KU139835.1	–	–	IV
Cinnamomum crenulicupulum	KU139838.1	–	–	IV
Cinnamomum curvifolium	KU139895.1	HM019397.1	–	IV
Cinnamomum cuspidatum	KU139839.1	–	–	IV
Cinnamomum daphnoides	MF110043.1	MF137962.1	–	IV
Cinnamomum doederleinii	KU139842.1	–	–	IV
Cinnamomum dubium	MH232442.1	MW408304.1	–	IV
Cinnamomum filipedicellatum	MH232443.1	–	–	IV
Cinnamomum foveolatum	MT628595.1	–	–	II
Cinnamomum glanduliferum	KX546415.1	MF137966.1	NC_057217.1	II
Cinnamomum goaense	MH232446.1	–	–	IV
Cinnamomum heyneanum	KU139847.1	MG209136.1	SY9391	IV
Cinnamomum insularimontanum	KY271510.1	AF268782.1	OL544942.1	IV
Cinnamomum javanicum	KU139852.1	–	–	IV
Cinnamomum jensenianum	KU139853.1	HM019391.1	–	IV
Cinnamomum kalbaricum	KU139844.1	–	–	IV
Cinnamomum keralaense	MH232450.1	–	–	IV
Cinnamomum kotoense	KY271518.1	KY296429.1	NC_050346.1	IV
Cinnamomum laubatii	KU139855.1	–	–	IV
Cinnamomum liangii	KU139856.1	–	–	IV
Cinnamomum litseifolium	MH232451.1	MW408307.1	–	IV
Cinnamomum longipaniculatum	KX546420.1	KY223702.1	MW553040.1	II
Cinnamomum longipetiolatum	KU139858.1	–	–	IV
Cinnamomum loureiroi	MF110051.1	MF137971.1	–	IV
Cinnamomum macrocarpum	MN519276.1	–	–	IV
Cinnamomum mairei	KU139859.1	–	–	IV
Cinnamomum malabatrum	MH232453.1	KY966336.1	–	IV
Cinnamomum micranthum	KU139860.1	KJ686734.1	NC_035802.1	II
Cinnamomum migao		–	NC_058709.1	II
Cinnamomum mohananianum	MH232466.1	–	–	IV
Cinnamomum mollifolium	KU139861.1	–	MW421302.1	II
Cinnamomum nilagiricum	MH232469.1	–	–	IV
Cinnamomum okinawense	KU139863.1	–	–	IV
Cinnamomum oliveri	KU139865.1	–	KT716496.1	IV
Cinnamomum osmophloeum	KY271528.1	KY296439.1	MT384386.1	IV
Cinnamomum paiei	MF110053.1	MF137973.1	–	IV
Cinnamomum parthenoxylon	KX546421.1	KX546120.1	MH050971.1	II
Cinnamomum pauciflorum	–	–	MW421303.1	IV
Cinnamomum perrottetii	MH232470.1	–	–	IV
Cinnamomum pingbienense	KU139873.1	–	–	IV
Cinnamomum pittosporoides	DQ124269.1	KX546125.1	NC_048978.1	IV
Cinnamomum platyphyllum	KU139875.1	HM019396.1	–	II
Cinnamomum polderi	MF110056.1	MF137976.1	–	IV
Cinnamomum propinquum	KU139876.1	–	–	IV
Cinnamomum racemosum	MF110044.1	MF137964.1	–	IV
Cinnamomum reticulatum	KU139879.1	MF623431.1	–	IV
Cinnamomum rhynchophyllum	KU139880.1	–	–	IV
Cinnamomum rigidissimum	KU139881.1	–	–	IV
Cinnamomum riparium	MH232473.1	MF072390.1	–	IV
Cinnamomum saxatile	KU139882.1	–	–	IV
Cinnamomum septentrionale	KU139883.1	–	MW421306.1	II
Cinnamomum sintoc	KU139884.1	–	–	IV
Cinnamomum subavenium	GU598528.1	KJ686740.1	MW801140.1	IV
Cinnamomum tamala	MF110058.1	MH232535.1	–	IV
Cinnamomum tavoyanum	MH232475.1	MF072387.1	–	IV
Cinnamomum tazia	KP092859.1	KX546123.1	–	IV
Cinnamomum tenuifolium	KU139892.1	HQ427110.1	–	II
Cinnamomum tenuipile	KU139893.1	KR533062.1	NC_057069.1	IV
Cinnamomum travancoricum	MH232479.1	MF547522.1	–	IV
Cinnamomum tsangii	KU139900.1	–	–	IV
Cinnamomum tsoi	KU139901.1	–	–	IV
Cinnamomum verum	KU139902.1	MH232541.1	NC_035236.1	IV
Cinnamomum walaiwarense	MH232490.1	–	–	IV
Cinnamomum wightii	MH232487.1	–	–	IV
Cinnamomum wilsonii	KU139904.1	KY223672.1	MW800949.1	IV
Cinnamomum yabunikkei	–	–	NC_044864.1	IV
Damburneya ambigens	KX509828.1	KX509888.1	–	I
Damburneya colorata	MK507234.1	MK507302.1	–	I
Damburneya coriacea	MF110063.1	MF137983.1	–	I
Damburneya gentlei	KX509830.1	KX509890.1	–	I
Damburneya guatemalensis	MF110015.1	MF137935.1	–	I
Damburneya inconspicua	MK507235.1	MK507303.1	–	I
Damburneya martinicensis	KX509831.1	KX509891.1	–	I
Damburneya minima	MK507236.1	MK507304.1	–	I
Damburneya parvissima	MK507237.1	MK507305.1	–	I
Damburneya patens	KX509832.1	KX509892.1	–	I
Damburneya purpurea	AF272293.1	EU153974.1	–	I
Damburneya salicifolia	AF272294.1	–	–	I
Damburneya smithii	MK507238.1	MK507306.1	–	I
Damburneya umbrosa	MK507239.1	MK507307.1	–	I
Dicypellium caryophyllaceum	MK507240.1	–	–	I
Dicypellium manausense	AF272270.1	AF268775.1	–	I
Endlicheria anomala	AF363371.1	–	–	I
Endlicheria bracteolata	AF363372.1	–	–	I
Endlicheria chalisea	MK507241.1	AF268756.1	–	I
Endlicheria citriodora	MK507242.1	AF268757.1	–	I
Endlicheria dysodantha	AF363373.1	–	–	I
Endlicheria glomerata	MF110064.1	MF137984.1	–	I
Endlicheria gracilis	AF363374.1	–	–	I
Endlicheria longicaudata	AF363375.1	MK507311.1	–	I
Endlicheria metallica	AF363376.1	–	–	I
Endlicheria multiflora	AF363377.1	MF786039.1	–	I
Endlicheria paniculata	AF363378.1	–	–	I
Endlicheria pyriformis	MF110066.1	MF137986.1	–	I
Endlicheria reflectens	AF272274.1	AF268758.1	–	I
Endlicheria rubra	AF363380.1	–	–	I
Endlicheria ruforamula	AF363381.1	–	–	I
Endlicheria sprucei	AF363382.1	–	–	I
Endlicheria szyszylowiczii	MF110067.1	MF137987.1	–	I
Endlicheria tessmannii	AF363383.1	–	–	I
Kubitzkia mezii	AF272276.1	AF268772.1	–	I
Kuloa ikonyokpe	AF272305.1	–	–	V
Kuloa usambarensis	MN431689.1	MN431714.1	–	V
Licaria armeniaca	MK507245.1	MK507314.1	–	I
Licaria bahiana	MF110068.1	MF137988.1	–	I
Licaria cannella	AF272280.1	AF268773.1	–	I
Licaria crassifolia	MF110069.1	MF137989.1	–	I
Licaria guianensis	AF272281.1	KX248791.1	–	I
Licaria martiniana	AF272279.1	KX248795.1	–	I
Licaria pachycarpa	MK507247.1	MK507316.1	–	I
Licaria rodriguesii	MK507248.1	MK507317.1	–	I
Licaria triandra	AF272282.1	AF268774.1	–	I
Machilus thunbergii	KX546512.1	–	NC_038204.1	Outgroup
Mespilodaphne cymbarum	MK507249.1	MK507318.1	–	I
Mespilodaphne quixos	MF110080.1	KX509937.1	OM135246.1	I
Mespilodaphne veraguensis	AF272319.1	–	–	I
Nectandra acutifolia	KX509834.1	KX509894.1	–	I
Nectandra amazonum	FM957816.1	KX509895.1	–	I
Nectandra angusta	KX509835.1	KX509896.1	–	I
Nectandra angustifolia	KF420965.1	KF421030.1	MF939340.1	I
Nectandra apiculata	KX509836.1	KX509897.1	–	I
Nectandra barbellata	KX509837.1	KX509898.1	–	I
Nectandra citrifolia	KX509842.1	–	–	I
Nectandra cuneatocordata	KX509843.1	KX509903.1	–	I
Nectandra cuspidata	AF272291.1	EU153966.1	–	I
Nectandra discolor	KX509844.1	KX509904.1	–	I
Nectandra grandiflora	KF420973.1	KF421022.1	–	I
Nectandra herrerae	KX509846.1	KX509906.1	–	I
Nectandra hihua	KX509847.1	KJ426843.1	–	I
Nectandra lanceolata	KF420966.1	KF421026.1	–	I
Nectandra latissima	KX509848.1	KX509909.1	–	I
Nectandra laurel	KX509849.1	KX509910.1	–	I
Nectandra lineata	KX509839.1	EU153970.1	–	I
Nectandra lineatifolia	KX509851.1	KX509912.1	–	I
Nectandra matthewsii	KX509840.1	KX509900.1	–	I
Nectandra maynensis	KX509853.1	KX509914.1	–	I
Nectandra micranthera	KX509855.1	KX509916.1	–	I
Nectandra microcarpa	KX509856.1	KX509917.1	–	I
Nectandra nitidula	KX509857.1	KX509918.1	–	I
Nectandra obtusata	KX509858.1	KX509919.1	–	I
Nectandra olida	KX509859.1	KX509920.1	–	I
Nectandra oppositifolia	KX509860.1	KX509921.1	–	I
Nectandra paranaensis	KX509861.1	KX509922.1	–	I
Nectandra paucinervia	KX509862.1	KX509923.1	–	I
Nectandra psammophila	MF110070.1	KX509924.1	–	I
Nectandra puberula	KX509863.1	KX509925.1	–	I
Nectandra pulverulenta	KX509864.1	KX509926.1	–	I
Nectandra reflexa	KX509865.1	KX509927.1	–	I
Nectandra turbacensis	AF272295.1	AF268768.1	–	I
Nectandra villosa	GQ480373.1	KX509928.1	–	I
Ocotea aciphylla	DQ787422.1	MF137994.1	–	I
Ocotea ambrensis	MN431690.1	MN431716.1	–	I
Ocotea arcuata	MK507250.1	MK507319.1	–	I
Ocotea atirrensis	MF110071.1	MF137995.1	–	I
Ocotea aurantiodora	MK507251.1	MK507320.1	–	I
Ocotea auriculiformis	MN431691.1	MN431717.1	–	I
Ocotea balanocarpa	MK507252.1	MK507321.1	–	I
Ocotea bicolor	GQ480375.1	–	–	I
Ocotea botrantha	KX509867.1	KX509930.1	–	I
Ocotea brachybotrya	GQ480376.1	–	–	I
Ocotea brenesii	MK507253.1	MK507322.1	–	I
Ocotea bullata	AF272298.1	AF268778.1	–	I
Ocotea caniflora	MK507254.1	MK507323.1	–	I
Ocotea catharinensis	KF420963.1	KF421033.1	–	I
Ocotea ceanothifolia	AF272299.1	KX248927.1	–	I
Ocotea ciliata	MF110072.1	MF137996.1	–	I
Ocotea comoriensis	MN431692.1	MN431718.1	–	I
Ocotea complicata	MK507256.1	MK507325.1	–	I
Ocotea congregata	MK507257.1	MK507326.1	–	I
Ocotea corymbosa	GQ480377.1	–	–	I
Ocotea cujumary	MK507258.1	MK507327.1	–	I
Ocotea cymosa	MN431693.1	MN431719.1	–	I
Ocotea daphnifolia	MK507259.1	MK507328.1	OM135247.1	I
Ocotea dentata	MK507260.1	MK507329.1	–	I
Ocotea diospyrifolia	GQ480379.1	–	–	I
Ocotea divaricata	MK507261.1	MK507330.1	–	I
Ocotea domatiata	MK507262.1	MK507331.1	–	I
Ocotea elegans	MF110073.1	MF137997.1	–	I
Ocotea fasciculata	MK507263.1	MK507332.1	–	I
Ocotea floribunda	KX509868.1	KX509931.1	–	I
Ocotea foetens	AF272300.1	MN431720.1	OM135248.1	I
Ocotea gabonensis	MF110075.1	MF138000.1	–	I
Ocotea glaucosericea	MK507264.1	MK507333.1	–	I
Ocotea glomerata	GQ480380.1	–	–	I
Ocotea grayi	MN431697.1	MN431724.1	–	I
Ocotea guatemalensis	MK507266.1	MK507335.1	–	I
Ocotea guianensis	AF272302.1	AF268761.1	OM135249.1	I
Ocotea heydeana	AF272304.1	–	–	I
Ocotea holdridgeana	MK507267.1	MK507337.1	–	I
Ocotea indecora	MF110076.1	MF138001.1	–	I
Ocotea insularis	MK507269.1	MK507339.1	–	I
Ocotea involuta	MN431698.1	MN431725.1	–	I
Ocotea javitensis	MK507270.1	MK507340.1	–	I
Ocotea kenyensis	MN114140.1	MN431726.1	–	I
Ocotea keriana	MK507271.1	MK507341.1	–	I
Ocotea laetevirens	MK507272.1	MK507342.1	–	I
Ocotea lancifolia	GQ480383.1	–	–	I
Ocotea laxa	MK507273.1	MK507343.1	–	I
Ocotea lentii	MK507274.1	MK507344.1	–	I
Ocotea leptobotra	MK507275.1	–	–	I
Ocotea leucoxylon	KX509852.1	KX509913.1	–	I
Ocotea longifolia	GQ480385.1	–	–	I
Ocotea longipes	MN431701.1	MN431728.1	–	I
Ocotea macrocarpa	MN431702.1	MN431729.1	–	I
Ocotea macrophylla	AF272303.1	MK507336.1	–	I
Ocotea magnilimba	KX509870.1	KX509932.1	–	I
Ocotea malcomberi	AF272307.1	AF268779.1	–	I
Ocotea mascarena	MN431703.1	MN431730.1	–	I
Ocotea meziana	MK507276.1	MK507345.1	–	I
Ocotea micans	MK507277.1	MK507346.1	–	I
Ocotea minarum	MK507278.1	MK507347.1	–	I
Ocotea montana	MK507279.1	MK507348.1	–	I
Ocotea nervosa	MN431704.1	MN431731.1	–	I
Ocotea nigra	AF272308.1	KX248946.1	–	I
Ocotea nitida	GQ480387.1	MK507349.1	–	I
Ocotea oblonga	MK507280.1	EU153984.1	–	I
Ocotea odorifera	KX509871.1	KF421038.1	OM135250.1	I
Ocotea pauciflora	MK507281.1	AF268764.1	–	I
Ocotea percoriacea	AF272311.1	MK507351.1	–	I
Ocotea perforata	MN431705.1	MN431732.1	–	I
Ocotea pluridomatiata	GQ480389.1	–	–	I
Ocotea pomaderroides	GQ480390.1	MK507352.1	–	I
Ocotea porosa	KF420956.1	KF421040.1	OM135251.1	I
Ocotea porphyria	MF110079.1	MF138004.1	–	I
Ocotea praetermissa	KX509872.1	KX509934.1	–	I
Ocotea puberula	KF420951.1	KF421045.1	–	I
Ocotea pulchella	KX509873.1	KX509935.1	–	I
Ocotea purpurea	KX509874.1	KX509936.1	–	I
Ocotea racemosa	MK507283.1	MK507354.1	–	I
Ocotea ramosissima	GQ480393.1	–	–	I
Ocotea rivularis	MK507284.1	MK507355.1	–	I
Ocotea salvadorensis	KX509875.1	KX509938.1	–	I
Ocotea sambiranensis	MN431707.1	MN431734.1	–	I
Ocotea sassafras	MK507285.1	MK507356.1	–	I
Ocotea schomburgkiana	AF272315.1	–	–	I
Ocotea sessiliflora	MN431708.1	MN431735.1	–	I
Ocotea silvestris	GQ480394.1	–	–	I
Ocotea sinuata	KX509876.1	KX509939.1	–	I
Ocotea spectabilis	MK507287.1	MK507358.1	–	I
Ocotea spixiana	AF272316.1	–	–	I
Ocotea tabacifolia	–	–	OM135252.1	I
Ocotea teleiandra	MK507288.1	MK507359.1	–	I
Ocotea tenera	MF110082.1	MF138006.1	–	I
Ocotea tessmannii	MK507290.1	MK507361.1	–	I
Ocotea thouvenotii	MN431709.1	MN431736.1	–	I
Ocotea tomentella	AF272317.1	AF268765.1	–	I
Ocotea trichantha	MN431710.1	MN431737.1	–	I
Ocotea trichophlebia	MN431711.1	MN431738.1	–	I
Ocotea tristis	AF272318.1	–	–	I
Ocotea valeriana	MK507292.1	MK507363.1	–	I
Ocotea velloziana	GQ480395.1	–	–	I
Ocotea venulosa	MN431712.1	MN431739.1	–	I
Ocotea whitei	MK507286.1	MK507357.1	–	I
Ocotea zahamenensis	MN431713.1	MN431740.1	–	I
Paraia bracteata	MK507293.1	MK507364.1	–	I
Persea americana	AF272322.1	–	NC_031189.1	Outgroup
Phoebe sheareri	FM957849.1	–	NC_031191.1	Outgroup
Pleurothyrium cinereum	AF272329.1	AF268769.1	–	I
Pleurothyrium cuneifolium	KX509879.1	KX509941.1	–	I
Pleurothyrium insigne	AF272330.1	–	–	I
Pleurothyrium poeppigii	KX509880.1	KX509942.1	–	I
Pleurothyrium trianae	MK507294.1	MK507365.1	–	I
Rhodostemonodaphne capixabensis	GQ480398.1	–	–	I
Rhodostemonodaphne crenaticupula	AF272331.1	AF268759.1	–	I
Rhodostemonodaphne kunthiana	AF363384.1	FJ038959.2	–	I
Rhodostemonodaphne negrensis	MK507295.1	MK507366.1	–	I
Rhodostemonodaphne parvifolia	AF363386.1	MK507367.1	–	I
Rhodostemonodaphne peneia	AF363387.1	–	–	I
Rhodostemonodaphne praeclara	AF272332.1	AF268760.1	–	I
Rhodostemonodaphne recurva	AF272333.1	–	–	I
Rhodostemonodaphne scandens	AF272334.1	–	–	I
Sassafras albidum	EF491214.1	AF268793.1	–	III
Sassafras randaiense	EF491212.1	EF491222.1	MW337246.1	III
Sassafras tzumu	AF272336.1	EF491220.1	NC_045268.1	III
Umbellularia californica	AF272337.1	AF268777.1	–	I
Urbanodendron bahiense	MK507296.1	MK507368.1	–	I
Urbanodendron verrucosum	MK507297.1	MK507369.1	–	I

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