Syzygium+grande


 * Sea Apple Tree****.** //For the Sea Apple Cucumber, see Pseudocolochirus voilaceus.//

“The clearest way into the Universe is through a forest wilderness.” – John Muir  Figure 1 (Left): Underside of a //Syzygium grande// crown at the Singapore Botanic Gardens. (Photograph by Tan WA Lorraine)

= **__ Introduction __** = toc //Syzygium grande //(Wight) Walp. is a species that belongs to the family Myrtaceae. In the Malesian biogeographical region, there are 35 genera of Myrtaceae and //Syzygium// is the most abundant in species 1]. This includes species like //Syzygium aromaticum,// the popular spice commonly known as clove. //Syzygium grande// can be found throughout the tropical Asia in a large variety of habitats 2,3]. This includes coastal forest, where it can be commonly found in Singapore. //Syzygium grande// is also cultivated in Singapore as a roadside ornamental tree 3]. The common name is Sea Apple, with other names such as //Jambu Laut// or //Jambu Air Laut// 4]. Capable of growing up to 30 m in height, [|one towering individual] in Sentosa has also been awarded the Heritage Tree status.

 Although it is Common in Singapore, //S. grande// is a species that is part of the natural heritage in Singapore 5]. This page aims to encourage an appreciation for the biology, value, and taxonomic history of //S. grande// as a part of our native flora.

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=__ Biology __=

__ **Distribution ** __
Native Distribution: Thailand, India, Sri Lanka, Burma, Indo-China, Malaysia, Singapore, Borneo 2]  Worldwide distribution: Figure 2: Global distribution of //Syzygium grande// from the Encyclopedia of Life 6]. //Syzygium grande// is an introduced species in the New World Tropics but has one native species in Hawaii 6,7]. According to the IUCN Red List, //S. grande// has been listed as one of its accepted synonyms, //Syzygium firmum. Syzygium firmum// is considered Vulnerable, but the range description only includes Sri Lanka and needs updating [8 ].

__ **Description ** __
Cultivated //S. grande// along a road side. ||
 * Large tree up to 30 m tall and 80 cm in diameter. Dense crown, dark and glossy with oblong shape 3]. || [[image:http://uforest.org/Images/s_grande1.jpg]]
 * The flowers are up to 3 cm across, white, and sessile 2]. Each flower has many stamens about 10 mm long and one 9–15.4 mm style exceeding them.

Up to 14 cm long inflorescences can be found at the base of a leaf stalk, or at the ends of its branches 3]. The flowers are also very strongly fragrant 4]. || Flowers of // S. grande // and the downturned leaves. Close-up of a flower by [|Photos of NUS Toddycats, ICCS, and LKCNHM] on Flickr || Ripe fruits of // S. grande. // ||
 * It produces fleshy fruits that are ellipsoid, about 4 cm long and 3 cm wide that are green when ripe 3,4]. Each fruit has a prominent 6mm diameter apical calyx rim and one seed 3]. || [[image:http://uforest.org/Images/s_grande5.jpg]]
 * It has simple, opposite, stalked leaves that are broadly elliptic, about 16 cm long and 9 cm wide 2,3]. The leaves are darker green above, lighter green below, and each has a down-turned leaf tip. Two intramarginal veins run parallel to the leaf margin, with 9–16 pairs of side veins 2]. || [[image:http://i460.photobucket.com/albums/qq328/lurabrwi/Sgrandearrow.png width="362" height="259"]]Senescent leaf showing the distinct intramarginal vein, edited from uforest.org

Leaves of a seedling showing the leaf arrangement. (Photograph by Tan WA Lorraine) || Trunk of an individual from the Singapore Botanic Gardens. (Photograph by Tan WA Lorraine) || There are two varieties of //S. grande//, //S. grande// var. //grande// and var. //parviflorum//. //Syzygium grande// var. //parviflorum// differs from the former with generally smaller flowers. It is only known from the [|type] locality and from Sabah.
 * The bark is pale-grey to pink-brown, becoming cracked and flaky with age 3]. The base of the trunk is fluted but are not truly [|buttressed] 4]. || [[image:http://i460.photobucket.com/albums/qq328/lurabrwi/Trunk.jpg width="313" height="416"]]

__ Ecology __
//Syzygium grande// can be found in habitats which include: bamboo forest, savannah, edges of evergreen forest and coastal forest 2,3]. It is a perennial which grows relatively quickly and generally flowers and fruits from March to May in Singapore, with exceptions 4]. As the seeds are common and germinate readily, the species helps with natural regeneration 9]. The fruits are eaten by bats which helps with their dispersal 9]. //Syzygium grande// is also the caterpillar host plant for butterflies species such as the Centaur Oakblue (//Arhopala centaurus nakula//) and the Plain Plushblue (//Flos apidanus saturatus//) 10,11]. With birds, a Thai study in Khao Yai National Park found that although //Syzygium// spp. (as //Eugenia//) trees represented only 3% of all large trees in 302 sample plots, they managed to garner 26% of the hornbill nests recorded 12]. Knowing the range and habit of //S. grande//, //S. grande// could be included in this observation and be beneficial to the local species of hornbill in Singapore. Another bird association that has been observed is with Red-breasted Parakeets (//Psittacula alexandri//) in Singapore, where they have been observed to eat the flowers of //S. grande//. media type="youtube" key="myiHZfp_o6s" width="560" height="315"

__ Uses __
The trunk is used as timber, a medium hard wood known as ‘Kelat’ 9]. It is also an ornamental tree that was planted as a roadside tree in Singapore to act as fire breaks from lalang fires 3]. Recently, to reduce the risk of branch falls 13], roadside //S. grande// trees have been replaced by a shorter species (up to 20 m) //Tristaniopsis whiteana//, or [|River Tristania]. More information on the ornamental uses and cultivation of //S. grande// can be found [|here].

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=**__ Taxonomy __**=

**__Classification __**
8]
 * <span style="font-family: Arial,sans-serif; font-size: 12pt;">Kingdom: Plantae
 * <span style="font-family: Arial,sans-serif; font-size: 12pt;">Phylum: Tracheophyta
 * <span style="font-family: Arial,sans-serif; font-size: 12pt;">Class: Magnoliopsida
 * <span style="font-family: Arial,sans-serif; font-size: 12pt;">Order: Myrtales
 * <span style="font-family: Arial,sans-serif; font-size: 12pt;">Family: Myrtaceae
 * <span style="font-family: Arial,sans-serif; font-size: 12pt;">Genus: //Syzygium//
 * <span style="font-family: Arial,sans-serif; font-size: 12pt;">Species: //Syzygium grande//

**__<span style="font-family: Arial,sans-serif; font-size: 12pt;">Origin of Name __**
<span style="font-family: Arial,sans-serif; font-size: 12pt; text-align: left;">Greek //syzygos//, joined, referring to the paired leaves of this genus; Latin //grandis//, great, referring to the large leaf and flower bud 3].

**__<span style="font-family: Arial,sans-serif; font-size: 12pt;">Name and Type __**
//<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium grande //<span style="font-family: Arial,sans-serif; font-size: 12pt;">(Wight) Walp. was originally described as //Eugenia grandis// Wight (1841) 14] as seen in Figure 3.

<span style="font-family: Arial,sans-serif; font-size: 12pt;">Figure 3: Description of //Eugenia grandis// Wight. Taken from the [|Biodiversity Heritage Library].

<span style="font-family: Arial,sans-serif; font-size: 12pt;">There was no specific holotype assigned in the original description, with the given information only listed as “//Mergui//—Griffith //Silhet//”. As such, a lectotype was designated by Byng //et al//. 15] with the details as follows: “—BURMA [Myanmar]. Mergui, //Griffith// s.n. (K! [barcode K000821429]; isolectotype P! [barcode P05208756])”.

<span style="font-family: Arial,sans-serif; font-size: 12pt;">The accepted name was given by Walpers in 1843 16].

__<span style="font-family: Arial,sans-serif; font-size: 12pt;">Synonyms __
<span style="font-family: Arial,sans-serif; font-size: 12pt; text-align: left;">The synonyms were taken from [|The Plant List ver 1.1]:
 * 1) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Eugenia cymosa //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> Roxb. (Illegitimate)
 * 2) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Eugenia grandis //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> Wight
 * 3) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Eugenia laosensis //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> Gagnep.
 * 4) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Eugenia laosensis //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> var. //<span style="font-family: Arial,sans-serif; font-size: 12pt;">quocensis //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> Gagnep.
 * 5) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Eugenia montana //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> Wight (Illegitimate)
 * 6) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Jambosa firma //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> Blume
 * 7) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Jambosa grandis //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> (Wight) Blume
 * 8) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium firmum //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> (Blume) Thwaites
 * 9) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium gadgilii //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> M.R.Almeida (Illegitimate)
 * 10) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium grande //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> var. //<span style="font-family: Arial,sans-serif; font-size: 12pt;">parviflorum //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> Chantaran. & J.Parn.
 * 11) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium laosense //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> (Gagnep.) Merr. & L.M.Perry
 * 12) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium laosense //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> var. //<span style="font-family: Arial,sans-serif; font-size: 12pt;">quocense //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> (Gagnep.) H.T.Chang & R.H.Miao
 * 13) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium megalophyllum //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> Merr. & L.M.Perry
 * 14) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium montanum //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> Thwaites & Hook.f.
 * 15) //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium tamilnadensis //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> Rathakr. & V.Chithra (Illegitimate)

<span style="font-family: Arial,sans-serif; font-size: 12pt;">Roxburgh described the species in 1832 as //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Eugenia cymosa //<span style="font-family: Arial,sans-serif; font-size: 12pt;">, before Wight 17]. This description was deemed illegitimate because the name //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Eugenia cymosa //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> was already used by Lamarck in 1789 for //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium cymosum //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> (Lam.) DC. 18]. This is why Wight’s description contains Roxburgh’s name for the species as //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Eugenia cymosa non //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> Lam..

** __<span style="font-family: Arial,sans-serif; font-size: 12pt;">//Eugenia-Syzygium// split __ **
//<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> Gaertn. 19] is the largest genus in the Myrtaceae and has about 1200 species of evergreen trees and shrubs in the Old World tropics and subtropics 3]. Separating species has been difficult due to few diagnostic characteristics and great variability 20]. There has also been much confusion because many //Syzygium// species were originally described under the Neotropical genus //Eugenia//. <span style="font-family: Arial,sans-serif; font-size: 12pt;">Merrill and Perry were the first to provide prominent modern research for the taxonomic revisions of //Eugenia// L. and //Syzygium// Gaertn. 21]. It was only more than 30 years later that this argument gained more ground, when Schmid provided strong floristic evidence to separate the two genera 22]. Even then, there was practical difficulty in telling apart the leaves, making the trend unpopular 4]. In molecular data, Gadek //et al//. were the first to use //<span style="font-family: Arial,sans-serif; font-size: 12pt;">matK //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> data to show the separation of the Asian //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> from the Neotropical //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Eugenia //<span style="font-family: Arial,sans-serif; font-size: 12pt;"> 23]. Their analysis showed that //Syzygium// is separate from the Myrtoidae clade which //Eugenia// belongs to, with bootstrap certainty of 52%. This, coupled with molecular and morphological data from many other studies, has led to the conclusion that the separation is fact 7]. <span style="font-family: Arial,sans-serif; font-size: 12pt;">As molecular analyses grew in frequency, Soh and Parnell were able to focus on //Syzygium// in greater detail 24]. In their study, they investigated the phylogenetic relationships of subgenera within the //Syzygium// genus. Through analysing DNA sequences (ETS and ITS) and leaf morphology, they were able to generate a strict consensus tree from eight of the Most Parsimonious Trees (MPTs) (Figure 4). This tree has higher bootstrap support than the analysis of the DNA evidence alone. However, the exact relationships between //S. grande// and the closely related species are still unclear due to the polytomy. <span style="font-family: Arial,sans-serif; font-size: 12pt;">//Syzygium cumini// is also a tree that can be found in tropical Asia, but it is an exotic that has been introduced in Singapore <span style="font-family: Arial,sans-serif; font-size: 16px;"> 25] <span style="font-family: Arial,sans-serif; font-size: 12pt;">. The other species have distributions restricted to mainland Southeast Asia 26]. <span style="font-family: Arial,sans-serif; font-size: 12pt;">Figure 4: Strict consensus tree of eight MPTs retrieved from the combined analysis of ETS, ITS and morphological data. Bootstrap values >50% are indicated above the branches. Bayesian posterior probabilities are indicated below the branches. Dashed line indicates alternative branching retrieved from Bayesian analysis. Adapted from Soh and Parnell 24].

__ **<span style="font-family: Arial,sans-serif; font-size: 12pt;">Future Work ** __
<span style="font-family: Arial,sans-serif; font-size: 12pt;">For future work in the //Syzygium// genus, Parnell //et al//. 7] predicts that there will be a lot of logistical difficulty. <span style="font-family: Arial,sans-serif; font-size: 16px;">There is currently insufficient data for such a species rich, widespread and poorly known genus. Molecular data has been limited on the reliability of genes and phylogenetically important morphological data (such as leaf anatomy 24]) are still being discovered. These limit understanding <span style="font-family: Arial,sans-serif; font-size: 12pt;">what drives diversification in //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium //<span style="font-family: Arial,sans-serif; font-size: 12pt;">and the different breeding mechanisms of the species. <span style="font-family: Arial,sans-serif; font-size: 12pt;">Their work also showed that it may be operationally unfeasible to adhere strictly to the concept of monophyly 7]. Variation patterns of //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium //<span style="font-family: Arial,sans-serif; font-size: 12pt;">in Southeast Asia differ from Australasia partly due to different breeding systems, making applying a uniform species concept difficult. From this arises the debate among taxonomists for what is "sufficient mass" to justify the possibility of splitting the very large //<span style="font-family: Arial,sans-serif; font-size: 12pt;">Syzygium //<span style="font-family: Arial,sans-serif; font-size: 12pt;">into smaller genera. As such, there is a lot of potential in looking into the genus and //S. grande// would be part of the scrutiny in years to come. Back to Top]

=__ References __= Back to Top]
 * 1) van Balgooy (1998) Malesian seed plants: Portraits of tree families. Volume 2. Rijksherbarium/Hortus Botanicus. 307 pp.
 * 2) Parnell J & Chantaranothai P (2002) Myrtaceae. In: Santisuk T & Larsen K (eds.) Flora of Thailand. Volume 7 Part 4. The Forest Herbarium, Royal Forest Department, Bangkok, Thailand. Pp. 778–914.
 * 3) Ashton PS (2011) Myrtaceae. In: Soepadmo E, Saw LG, Chung RCK & Kiew R (eds.) Tree Flora of Sabah and Sarawak. Volume 7. Forest Research Institute Malaysia. Pp. 87–330.
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 * 7) <span style="background-color: #ffffff; color: #222222; font-family: Arial,sans-serif;">Parnell JAN, Craven LA & Biffin E (2006) Matters of Scale: Dealing with One of the Largest Genera of Angiosperms. <span style="background-color: #ffffff; color: #222222; font-family: Arial,sans-serif;">In: Hodkinson TR & Parnell JA (eds.) Reconstructing the tree of life: taxonomy and systematics of species rich taxa<span style="background-color: #ffffff; color: #222222; font-family: Arial,sans-serif;">. CRC Press, Boca Raton, Florida. Pp. 251–273.
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