Research Article |
Corresponding author: Fabian Brambach ( fbramba@gwdg.de ) Academic editor: Peter de Lange
© 2017 Fabian Brambach, James W. Byng, Heike Culmsee.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Brambach F, Byng JW, Culmsee H (2017) Five new species of Syzygium (Myrtaceae) from Sulawesi, Indonesia. PhytoKeys 81: 47-78. https://doi.org/10.3897/phytokeys.81.13488
|
Following ongoing ecological research on the tree diversity of the Indonesian island of Sulawesi, we describe five new species of Syzygium. These are the first descriptions of Syzygium species from the island since
Indonesia, Lore Lindu National Park, Myrtaceae , Sulawesi, Syzygieae , Syzygium , taxonomy, Wallacea
The botanical diversity of the Indonesian island of Sulawesi is poorly known and remains one of the least studied in Southeast Asia (de
Syzygium is the most - species rich genus of woody plants in Southeast Asia with around 1000 species but little is known of the genus in Wallacea, the biogeographically important transition zone between the Asian and Australian continental areas. As in the other Wallacean regions, the Maluku Islands and Lesser Sunda Islands, the Syzygium species of Sulawesi have never been revised or monographed so there is no robust baseline data of which species occur in the region. The last Syzygium species to be described from Sulawesi were by
Species of Syzygium are present in virtually all ecosystems of Sulawesi, and are often important components of the biological communities (van
In 2006–2007 and 2011–2012, the University of Göttingen, Germany, and Tadulako University, Palu, Indonesia conducted ecological fieldwork campaigns in Lore Lindu National Park (LLNP), Central Sulawesi. Difficulties in the identification of Syzygium specimens collected during these surveys motivated us to take a closer look at the taxonomy of the genus. Fortunately, the area of LLNP had been visited before by other botanists (
The specimens collected during our ecological fieldwork in LLNP (HC, 2006–2007; HC and FB, 2011–2012) were the starting point for this study. Duplicates of relevant specimens, including types, were deposited in L and the Indonesian herbaria BO and CEB (herbarium acronyms follow Thiers continuously updated). To identify our specimens, all Syzygium specimens from Sulawesi at A, B, BM, BO, E, GH, K, L, M and U were examined and all matching specimens sorted into morphospecies. We then attempted to identify our morphospecies using keys and floristic treatments from regions around Sulawesi: the Malay Peninsula (
Photographs in the field were taken using a Canon EOS 500D camera with a Tamron AF 18–200mm f/6.2–38 lens, for later photographs of dried material we used the same camera with a Tamron SP 90mm F/2.8 MACRO lens. Colours of dried specimens were compared to Munsell Soil-Color Charts (
Wood density (oven-dry mass per fresh volume) was determined from wood cores extracted with increment borers. The samples’ fresh volume was measured by Archimedes’ principle and weight was noted from the same samples after oven-drying for 48h at 105°C.
For the descriptions, flowers and fruits were boiled in dilute detergent for 5 minutes and dissected thereafter. Dimensions were measured using a ruler with 0.5 mm accuracy. All colours and measures given refer to dried and pressed material unless otherwise stated. We measured the distance of intramarginal veins from leaf margin in the proximal 2/3 of the blade; it usually decreases towards the apex. Likewise, we measured the distance of secondary veins in the central ½ of the leaf; it decreases near the base. Dimensions of flower buds are given including the anthopodium, those of the hypanthium excluding the anthopodium (if present).
Terminology for organs in Syzygium has been varied and often confusing, with authors using different terms for the same structures or similar terms for different structures.
We here adopt the detailed concepts of
Since several specimens found in herbaria contained very limited information about the respective collecting localities, we interpreted the locality data of all specimens cited in this paper and translated it into a common format. The format contains approximate coordinates in WGS 84 (if not given on the label), the nearest village, and the administrative divisions in descending order: Province, Kabupaten (Kab., Regency), and Kecamatan (Kec., District).
Specimens collected in Sulawesi by the Forest Research Institute Buitenzorg (Bogor), also called Boschproefstation or Boschbouwproefstation (
In the diagnoses, we give floral formulas for each species, following the format and recommendations of
Under Distribution and Habitat, we characterised the forest stands of the species which were found primarily in our (FB and HC) inventory plots by mentioning the families with the five highest family importance values (FIV). The FIV is an objective measure of importance of a family in a stand taking into account the number of individuals, number of species, and basal area of that family and comparing them to the stand total (see
We used GeoCAT (
All species here described are glabrous in all parts and possess flower characters placing them in the broadly defined Syzygium subg. Syzygium (
“Eugenia spec. BB“ (
„Myrtaceae sp. 10“ p.p. (
Syzygium balgooyi is characterised by long, elongate-clavate flowers, a character otherwise only known from the morphologically similar Syzygium schumannianum (Nied.)
INDONESIA. South Sulawesi (Sulawesi Selatan), Kab. Luwu Timur, Kec. Nuha, Between Soroako and Nickel plant site, c. 2°33'S, 121°22'E, 500 m, 10 Jul 1979: van Balgooy 3956 (flowers; holotype L [L.2517558]! [spirit collection L 0771145] [wood sample L 0708624], isotype A [A01143212]!).
Trees, up to 37 m tall, diameter at breast height ≤ 65 cm, trunk straight, ≤ 20 m tall, often fluted and with buttresses ≤ 3 m tall and 1 m out. Outer bark pale brown to bright red, peeling off in small or large sheets, inner bark dark red, usually paler towards inside, sometimes with little watery red sap, wood very hard and heavy, sapwood cream, clearly separated from the dark reddish brown heartwood. Young branchlets 1–2 × 1.5–4 mm, strongly flattened, the flat sides usually with two lateral, rounded ridges leading to the petioles and one central ridge continuing into the next internode, often resinous when dry, epidermis green, drying dusky red to reddish black and usually smooth; becoming terete, bark drying red to dark reddish brown, finely flaking and with conspicuous flaking remnants of epidermis.
Leaves (sub-)opposite. Petioles 2–12 × 1–3.5 mm, flat and sometimes narrowly winged above, rounded or keeled beneath, drying reddish black and smooth. Blades (4–) 7–11.5 (–16) × (1.5–) 3–5 (–9) cm, ratio (1.2–) 1.8–2.7 (–5), (narrowly) elliptic, obovate, or oblanceolate, base cuneate and attenuate at the very base or obtuse to rounded, apex usually rounded or obtuse, sometimes emarginate or acute, margin slightly to strongly revolute; (thick-)coriaceous, purple, pink, or reddish when young, fresh to dark glossy green above, paler glossy green beneath, drying dull to shiny, often resinous after drying, reddish brown to reddish black above, reddish brown to very dusky red beneath. Midrib channelled above, prominent and rounded or keeled, drying reddish black and smooth beneath. Secondary vein pairs (9–) 11–14 (–16), 4–12 (–15) mm apart, ± faint and lighter red than the lamina above, ± prominent and darker than the lamina beneath; intersecondary veins present. Tertiary veins sup-parallel near the midrib, reticulate towards the margins, ± faint above, faint or prominulous and darker than the lamina beneath. Inner intramarginal vein 1–5 mm from the leaf margin, ± looping; outer intramarginal vein < 1 mm from the leaf margin, often seemingly absent from leaf margin.
Inflorescences terminal and often in axils of distal leaf pair, rather dense panicles, 5–10 cm long, peduncles 1–6 cm long, axes subangular or rounded, flattened, resinous after drying. Bracts c. 3 mm long, linear, pellucid-dotted, caducous; bracteoles 2 per flower, sometimes seemingly 4 (by contraction of the ultimate inflorescence axes?), 1 mm long.
Flowers 5–15 per inflorescence, within the panicles in monads or clusters of 2–4, 4-merous, anthopodium absent, c. 20–30 mm in diameter at anthesis, mature buds 20–30 × 3–6 mm. Hypanthium 20–30 × 5–7 mm, elongate-clavate, yellowish green, drying smooth black, hypanthium rim 15 mm long, glandular inside. Calyx lobes c. 2 × 2 mm, claw- or hood-shaped. Petals c. 4 × 3 mm, ± obovate, pale green. Stamens c. 100, filaments 10–20 mm long, pale green, anthers c. 0.5–0.8 mm long, ellipsoid, yellow. Ovary bilocular, locules surrounded by spongy tissue, ovules c. 15–20 per locule, ascending, ± arranged in 2 longitudinal rows. Style 25–35 mm long, pointed.
Fruits 1–2-seeded, 27–33 × 12–16 mm, ampulliform, yellowish green (immature?), drying black, smooth or slightly warty, pericarp c. 1 mm thick, leathery when fresh, ± woody when dried, hypanthium rim 8–12 mm long, 4–5 mm in diameter.
Seeds 13–15 × 9–10 mm, ellipsoid, testa cartilaginous, attached to the pericarp, cotyledons free from the testa, ± half-globose, minutely verrucose, facing surfaces undulate.
The species is named after Max Michael Josephus van Balgooy (*1932), botanist and authority on Southeast Asian plant taxonomy, identification, and biogeography. He collected over 900 specimens during a Dutch-Indonesian expedition to Sulawesi in 1979, among them the type specimen of this species. We enjoyed the privilege of learning from Max during several stays at the herbarium in Leiden and receiving his help with the identification of our specimens collected in Central Sulawesi.
Flowering specimens have been encountered throughout the year without any apparent association with geography or climate. Fruiting specimens have been recorded in May (de Vogel 5413) and September (sight record by FB).
Syzygium balgooyi is restricted to Sulawesi and widespread across the island (Figure
The AOO of 64 km² would place Syzygium balgooyi in the category “Endangered” (EN), despite its wide distribution in Sulawesi (Figure
Cenke hutan (= forest clove, Indonesian, de Vogel 2651), Jambu (general name for Syzygium, Indonesian, NIFS bb 33081), Rokobako (NIFS Cel./II-385), Tambeanitu (Bahasa Behoa, Brambach et al. 1047, 1083, 1290, 1316), Wawahuling (Bahasa Tondano, Koorders 18251, see
Among Syzygium species of Sulawesi, S. balgooyi can be recognised in the field by its tall stature (Figure
Leaf size and thickness are quite variable (Figure
Syzygium balgooyi and S. schumannianum are difficult to separate in vegetative state. S. balgooyi usually has leaves with rounded, obtuse, emarginate, or acute tips, whereas they are shortly acuminate in S. schumannianum, but there are exceptions in both species. Flowers and fruits of the two species also share the same structure but there are two important differences which we consider sufficient to warrant specific separation: Firstly, as indicated by the original name Eugenia neurocalyx Schumann nom. illeg. (in
The wood of S. balgooyi is used for construction in North Sulawesi, but is not water-resistant (
(Paratypes). INDONESIA. North Sulawesi (Sulawesi Utara): Kab. Minahasa, Kec. Kakas, Old-growth forest Pinamorongan, c. 1°08'N, 124°56'E (“Noord-Celebes, Residentie Menado, Pinamorongangebergte bij Kakas”), 500 m, 30 Jan 1895: Koorders 18251 (sterile; L [L.2517502]! [L.2535743]!).
Kota Bitung, Kec. Ranuwulu, southern part of Wiau Forest Reserve (Hutan Lindung G. Wiau), base of Mt Klabat, c. 1°28'N, 125°03'E, 400 m, 1 Nov 1973: de Vogel EF 2651 (flowers; L [L.2535729]! [L.2535730]! [wood sample L 0204047]).
Central Sulawesi (Sulawesi Tengah), LLNP: Kab. Poso, Kec. Lore Utara, west slope of Mt Rorekautimbu, c. 1°16'S, 120°16'E, 1700 m, 15 May 1979: van Balgooy MMJ 3371 (sterile; L [L.2535697]!).
Kab. Poso, Kec. Lore Utara, west slope of Mt Rorekautimbu, c. 1°16'S, 120°17'E, 2000 m: 5 May 1979: Tantra IJM 1589 (sterile; L [L.2517457]!), & 1592 (sterile; L [L.2535672]!); ibid. loco, 17 May 1979: de Vogel EF 5413 (fruits; BO [BO-1686561], K [K001024419]!, L [L.2517562]! [L.2517563]!, [wood sample L 0708565]).
Kab. Poso, Kec. Lore Utara, 4 km E of Wuasa, c. 200 m N of Rumuku waterfall, tree-inventory plot Torongkilo, 1°24.9'S, 120°16.7'E, 1450 m, 6 Mar 2012: Brambach F, Mangopo H, Firdaus, Faber M, Tiranda R 1478 (sterile; BO [BO-1938440]!, CEB, L!) & 1564 (flower buds; BO [BO-1938441]!, CEB, K [K000993483]!) & 1583 (sterile; GOET [GOET020022]!).
Kab. Poso, Kec. Lore Tengah, 9 km NW of Bariri, 100 m E of climate tower, tree-inventory plot Bariri NE, 1°39.4'S, 120°10.5'E, 1400 m: Jul 2007: Culmsee H y896 (sterile; CEB, L!); ibid. loco, 21 Aug 2011: Brambach F, Mangopo H, Firdaus, Faber M, Tiranda R 0861 (sterile; BO [BO-1938438]!, CEB, GOET [GOET020025]!) & 0889 (sterile; BO [BO-1938439]!, CEB, L!) & 0907 (sterile; CEB, GOET [GOET020024]!, L!).
Kab. Poso, Kec. Lore Tengah, 9 km NW of Bariri, 80 m south of climate tower, tree-inventory plot Bariri S, 1°39.5'S, 120°10.4'E, 1400 m, Jul 2007: Culmsee H 1459 (sterile; CEB, GOET [GOET020006]!) & 1495 (sterile; BO [BO-1938457]!, CEB); ibid loco, Jul 2007: Culmsee H r808 (sterile; CEB, GOET [GOET020008]!).
Kab. Poso, Kec. Lore Tengah, 7 km WNW of Hanggira, E flank of Mt Dali, tree-inventory plot Pantakleabae, 1°42.0'S, 120°09.0'E, 1950 m: 3 Mar 2011: Culmsee H, Brambach F, Mangopo H, Firdaus, Faber M, Tiranda R r2162 (sterile; CEB, GOET [GOET020021]!) & r2254 (sterile; BO [BO-1927087], CEB, GOET [GOET020023]!); ibid. loco, 30 Mar 2011: Brambach F, Mangopo H, Firdaus, Faber M, Tiranda R 0038 (sterile; BO [BO-1926965], CEB, GOET [GOET020027]!, K [K000993482]!, L!) & 0058 (sterile; BO [BO-1926969]!, [BO-1926970]!, CEB, GOET [GOET020033]!) & 0082 (sterile; BO [BO-1938382]!, CEB, GOET [GOET020030]!, L!) & 0097 (sterile; CEB, GOET [GOET020029]!, L!); ibid. loco., 23 Jan 2012: Brambach F, Mangopo H, Firdaus, Faber M, Tiranda R 1333 (sterile; CEB, GOET [GOET020020]!, L!).
Kab. Sigi, Kec. Kulawi, 2.4 km ENE of Toro, NE edge of Pono Valley, tree-inventory plot Pono, 1°29.7'S, 120°03.4'E, 1050 m: 4 Aug 2006: Culmsee 125 (sterile; BO [BO-1938456]!, CEB, L!) & 209 (sterile; CEB, K [K000993486]!); ibid. loco, Jul 2007: Culmsee r211 (sterile; CEB, GOET [GOET020009]!).
Kab. Sigi, Kec. Kulawi Selatan, 4 km E of Watukilo, following footpath to Mt Tokepangana, tree-inventory plot Tokepangana, 1°36.9'S, 120°04.4'E, 850 m, 16 Apr 2011: Brambach F, Mangopo H, Firdaus, Faber M, Tiranda R 0176 (sterile; BO [BO-1926967]!, CEB, GOET [GOET020028]!, L!) & 0206 (sterile; BO [BO-1926968]!, CEB) & 0283 (sterile; BO [BO-1926973]! [BO-1926974]!, CEB, GOET [GOET020032]!, K [K000993481]!, L!) & 0319 (BO [BO-1926934]!, CEB) & 0332 (BO [BO-1926966]!, CEB) & 0363 (BO [BO-1926919]!, CEB).
Kab. Sigi, Kec. Kulawi Selatan, 4 km ENE of Watukilo, 400 m N of Mboe River, tree-inventory plot Rantena, 1°36.2'S, 120°04.5'E, 700 m: 17 Jun 2011: Brambach F, Mangopo H, Firdaus, Faber M, Tiranda R 0466 (sterile; BO [BO-1938383]!, CEB, GOET [GOET020031]!); ibid. loco, 21 Jun 2011: Brambach F, Mangopo H, Firdaus, Faber M, Tiranda R 0628 (sterile; CEB, GOET [GOET020026]!, L!).
Kab. Sigi, Kec. Nokilalaki, 4.3 km SSW of Tongoa, NW flank of Mt Nokilalaki, ca. 400 m S of Shelter 2, tree-inventory plot Nokilalaki 2, 1°14.6'S, 120°09.1'E, 1850 m, Sep 2007: Culmsee 2923 (sterile; CEB, L) & 3075 (sterile; BO [BO-1938463]!, CEB).
Kab. Sigi, Kec. Nokilalaki, 4.3 km SSW of Tongoa, NW flank of Mt Nokilalaki, ca. 500 m SSE of Shelter 2, tree-inventory plot Nokilalaki 1, 1°14.7'S, 120°09.2'E, 1900 m, Aug 2007: Culmsee 2636 (sterile; CEB, L!) & 2641 (sterile; BO [BO-1938462]!, CEB, GOET [GOET020007]!) & 2721 (sterile; CEB, K [K000993487]!).
Kab. Tojo Una-una, Kec. Ulubongka. N slope of Mt Katopas, 1°9.8'S, 121°26.9'E, 1100 m, 4 Sep 2014: Sight record by F Brambach (photograph Figure
West Sulawesi (Sulawesi Barat): Kab. Mamasa. Kec. Mamasa, near Osango c. 2°56'S, 119°19'E (“Celebes en Ond. Boven Binoeang, ca. Osango”), c. 1500 m, 1 Jul 1939: Netherland's Indies Forest Service (NIFS) bb 28293 (sterile; L [L.2529832]!).
South Sulawesi (Sulawesi Selatan): Kab. Luwu, Kec. Ponrang, near Kampung Tampa, c. 3°11'S, 120°13'E (“Celebes en Ond. Palopo, Bakka, Kampoeng Tampa”), c. 100 m, 15 Sep 1941: NIFS bb 33081 (flowers; L [L.2535805]!).
Kab. Luwu Timur: Kec. Malili, Ussu, c. 2°36'S, 121°06'E (“Selebes, Malili, Oesoe): c. 300 m, 13 Jul 1931: NIFS Cel./II-385 (flower buds; L [L.2535679]!); ibid. loco, c. 400 m, 19 Jun 1934: NIFS Cel./II-293 (sterile; L [L.2517541]!); ibid. loco, 100 m, 28 Mar 1941: NIFS bb 32595 (sterile; BO [BO-1304600], L [L.2517463]!).
Kab. Luwu Timur, Kec. Wasuponda, Larona, c. 2°45'S, 121°20'E, 500–1000 m (“Celebes. Goud. Celebes, Ond. afd. Malili, nabij La Rona”), n.d.: NIFS bb 1843 (sterile; L [L.2535842]!) & bb 1895 (sterile; L [L.2535843]!).
Kab. Luwu Timur, Kec. Nuha, Hills W of Soroako, c. 2°31'S, 121°19'E, 550 m, 17 Jun 1979: van Balgooy MMJ 3767 (old inflorescences; L [L.2535910]! [wood sample L 0708626]).
Morphological characters of Syzygium balgooyi and S. schumannianum. Syzygium balgooyi (a–k): a c. 20 m tall trunk b upper leaf surface; c lower leaf surface d bark with slash e ridged shoot apex with subopposite leaves f flowers at different stages during anthesis and fruit g trunk base with steep narrow buttresses h inflorescence with flower buds i shoot with young leaves j dried flowers before and during anthesis and longitudinal section of flower k dried fruit and longitudinal section of fruit showing two cotyledons. Syzygium schumannianum (l–m): l dried flowers before and during anthesis m dried fruit. a–b and h–iBrambach et al. 1564cBrambach et al. 0861d–eBrambach et al. 0628f sighting on Mt Katopas by FB gBrambach et al. 0889; j holotype van Balgooy 3956 [L.2517558] kde Vogel 5413 [L.2517563] lWiakabu et al. LAE 50571 [L.2535534] mBrass 13610 [L.2524420]. All scale bars: 1 cm.
“Myrtaceae sp. 9” (
Syzygium contiguum is a species of treelets with slender, angular young branchlets and (sub-)sessile, chartaceous leaves with few (8–13), distinct secondary veins, two marginal veins, and conspicuous cordate bases; the basal lobes of opposed leaves often reach each other. The dense or lax paniculate inflorescences are terminal or arise from the upper leaf axils and bear small (5–6 × 3–4 mm in mature buds) pyriform flowers with numerous white stamens. The species is similar to Syzygium urdanetense (Elmer)
INDONESIA. Central Sulawesi (Sulawesi Tengah), LLNP, Kab. Sigi, Kec. Kulawi, 2.4 km ENE of Toro, NW of Pono Valley, tree-inventory plot Pono, 1°29.7'S, 120°03.4'E, 1050 m, Jul 2006: Culmsee H 535 (flowers; holotype L[L.3962133]!, isotype CEB).
Treelets, up to 10 m tall, diameter at breast height ≤ 11 cm. Bark and wood not known. Young branchlets 0.5–1 × 1–2 mm, slender, rectangular in cross section, sometimes narrowly winged, epidermis drying dark reddish brown, smooth; soon becoming terete with 4 ridges and eventually terete, bark pale or yellowish brown with flaking remnants of epidermis; with (1–) 2 (–4) pairs of ≤ 2 mm long, caducous cataphylls near the base of the current flush.
Leaves opposite, (sub-)sessile. Petioles 0–1.5 × 1–2 mm, absent or very short and stout, drying very dusky red. Blades (6.5–) 9–14 (–19) × (2.3–) 3.5–5 (–6.1) cm, ratio (1.9–) 2.5–3.2 (–3.6), narrowly elliptic or lanceolate, rarely oblanceolate, base distinctly cordate (or auriculate), basal lobes of opposed leaves often touching each other, apex (long-)acuminate or caudate, margin flat or sometimes minutely revolute; chartaceous, drying dull to satin, dark reddish brown to very dusky red above, (dark) reddish brown beneath; sometimes with scattered black gland dots. Midrib channelled above, prominent, rounded, and darker than the lamina beneath. Secondary vein pairs 8–12, (3–) 5–11 (–18) mm apart, slightly sunken or sometimes slightly prominent, rather inconspicuous above, very prominent and darker than the lamina beneath; some intersecondary veins usually present. Tertiary veins sub-parallel near the midrib to reticulate towards the margin, faint above, prominulous beneath. Inner intramarginal vein 3–7 mm from leaf margin, hardly looping; outer intramarginal vein 0.5–2 mm from leaf margin.
Inflorescences terminal and in the axils of 1–2 distal leaf-pairs, ± lax panicles, (2.5–) 3.5–7.5 (–11) cm long, peduncles 1–3.5 cm long, axes (sub-)angular, flattened. Bracts c. 0.5–2 (–7) mm long, lowermost foliaceous, caducous, others deltate, keeled, ± persistent; bracteoles 2 per flower, 0.5–1 mm long, similar to bracts.
Flowers ≤ 40 per inflorescence, within the panicles in monads or triads, 4-merous, anthopodium absent, c. 15 mm in diameter at anthesis, mature buds 5–6 × 3–4 mm. Hypanthium 4–5 × 3–5.5 mm, obconical to infundibuliform, gland-dotted or ± smooth, hypanthium rim 2 mm long. Calyx lobes 0.5–1 × 1–2.5 mm, deltate first, becoming broadly rounded and eventually splitting irregularly at anthesis. Petals 3–6 × 3–6 mm, pseudocalyptrate, orbicular, gland-dotted. Stamens c. 80–100, filaments 6–10 mm long, white, anthers c. 0.5 mm long, ellipsoid. Ovary bilocular, locules subtended by spongy tissue, ovules c. 8 per locule, spreading. Style 6–8 mm long, pointed.
Fruits 2-seeded, 1.1–1.3 × 1.8–1.9 cm, globose to oblate, drying smooth, pericarp c. 2 mm thick, hypanthium rim c. 5 mm in diameter.
Seeds 9–10 × 12–13 mm, half-moon shaped.
The specific epithet refers to the leaf bases of opposing leaves which, due to their cordate shape and the short petioles, often approach or touch each other.
In Central Sulawesi a slight dry season usually lasts from May to September or October. Flowering was observed during the wet and dry seasons: in July 2016, January/February 2007, July 2007 in Pono and in April 1975 on Mt Nokilalaki.
According to our present knowledge, the species is endemic to the province of Central Sulawesi. It has been recorded from only three localities in and around LLNP at 1000–1150 m elevation (Figure
In the Pono inventory plot, the species was found in undisturbed submontane rainforest on flat terraces with Sideralic Cambisols (
Syzygium contiguum has a limited geographical distribution (estimated EOO 557 km²) and seems to be restricted to submontane forest within a narrow elevational belt. We assume that the estimated AOO of 12 km² is unrealistically low, due to limited collection activities in Central Sulawesi. However, only the collection locality of Meijer 9572 seems to be covered by intact forest habitat. The other two localities are small forest fragments (Widjaja EAW 3502) and forest with recent deforestation activities in close proximity (Pono inventory plots, detected using the Global Forest Change website,
Syzygium urdanetense (as Eugenia urdanetensis,
Two fruiting specimens collected at low elevations (200–300 m) on Sulawesi’s Southeast Peninsula, Prawiroatmodjo & Maskuri 1231 [L.2517450] and 1957 [L.2517547], are morphologically similar to S. contiguum as defined above except for the leaf tips which are not long-acuminate. In the absence of flowering material, and because of the different habitat and distribution, we prefer not to include them here at present, but future additional collections may prove otherwise.
We choose Culmsee 535 as type specimen because it contains flowers in all stages of maturity although unfortunately, it was collected with only two duplicates (in CEB and L). Nevertheless, the more widely distributed paratypes collected by HC at the type locality all belong to the same population as the type.
(Paratypes). INDONESIA. Central Sulawesi (Sulawesi Tengah), LLNP: Kab. Poso, Kec. Nokilalaki, N slopes of Mt Nokilalaki. (“Celebes, central part, area of Mt. Nokilalaki, Loro Kalimata Reserve”), 1°13'S, 120°08'E, ± 1000 m, 24 Apr 1975: Meijer 9572 (flowers; L [L.2535817]!, US [US-2995269] photo!).
Kab. Poso, Kec. Lore Peore, Road to Napu from camp Dongi-dongi, 1°31.2'S, 120°22.4'E, 1127 m, 26 Dec 1988: Widjaja EA EAW 3502 (fruits; BO [BO-1917489]! [BO-1917490]!, K! , L).
Kab. Sigi, Kec. Kulawi, 2.4 km ENE of Toro, NW of Pono Valley, tree-inventory plot Pono, 1°29.7'S, 120°03.4'E, 1050 m, Jul 2006: Culmsee H 284 (flowers; GOET [GOET020010]!) .
Kab. Sigi, Kec. Kulawi, 2.4 km ENE of Toro, NW of Pono Valley, tree-inventory plot “Pono”, 1°29.7'S, 120°03.4'E, 1050 m, Jan 2007: Culmsee H y410 (flower buds; BO [BO-1938450]! [BO-1938451]!, CEB, GOET [GOET020012]!, K [K000993488]!, L!); ibid. loco, Jul 2007: Culmsee H r463 (flower buds; BO [BO-1938464]!, CEB, GOET [GOET020011]!, K [K000993489]!, L!!) & y503 (flower buds; CEB, GOET [GOET020013]!) & y514 (flower buds; BO [BO-1938452]!, CEB) & y581 (flower buds; CEB, L!) & y582 (flower buds; CEB, K [K000993490]!) & y592 (flower buds; BO [BO-1938453]! [BO-1938454]!, CEB) & y595 (flower buds; CEB, GOET [GOET020014]!, L!).
„Myrtaceae sp. 10“ p.p. (
Syzygium devogelii is a species of treelets characterised by slender, narrowly winged young branchlets, medium-sized narrowly elliptic leaves, straight and distinct secondary veins connected by an intramarginal vein impressed above and prominent beneath, small flowers (5 × 3 mm in bud) in terminal inflorescences that develop into rather large fruits (c. 20 × 25 mm), mature seeds lacking a testa, and cotyledons with echinate outer surfaces. The species is morphologically similar to Syzygium perspicuinervium (Merr.)
INDONESIA. Central Sulawesi (Sulawesi Tengah), LLNP, Kab. Poso, Kec. Lore Utara, west slope of Mt Rorekautimbu, c. 1°17.5'S 120°16.3'E, 1350 m, 11 May 1979: de Vogel EF 5293 (fruits; holotype L [L.2535665]! [L.2535666]!; isotype K!).
Trees, up to 13 m tall, diameter at breast height ≤ 13 cm, trunk ≤ 7 m tall. Outer bark whitish to brown, mealy or peeling off in thin sheets, inner bark pale or dark red, wood cream-coloured. Young branchlets 1–2.5 × 2–3 mm, ± flattened, angular or oblong in cross section with 4 narrow wings, epidermis dark red when young, drying reddish or yellowish brown, smooth; becoming rounded with 4 ridges, bark (yellowish) brown, peeling off in thin sheets.
Leaves (sub-)opposite. Petioles 7–16 × 1–3 mm, channelled above, rounded beneath, epidermis drying smooth or with transverse cracks. Blades (12.5–) 14–19 (–22.5) × (4–) 4.5–7 (–8.5) cm, ratio (2.1–) 2.6–3.3 (–4), narrowly elliptic (or lanceolate), base cuneate or obtuse, apex acuminate, margin revolute; chartaceous or coriaceous, red or pink when young, above, beneath, drying dull to satin, variable in colour from greyish brown and olive grey to very dusky red above, dull to satin and, dark reddish brown beneath; pellucid dots rather few, visible or not on both sides. Midrib channelled above, very prominent, rounded, smooth and drying darker than the lamina beneath. Secondary vein pairs (9–) 11–14 (–17), 5–22 mm apart, channelled or impressed above, prominent and drying darker than the lamina beneath, straight or slightly arching from the midrib; intersecondary veins sometimes present. Tertiary veins dense, ± ladder-like and perpendicular to the midrib, faint above, prominulous beneath. Inner intramarginal vein 2–9 mm from leaf margin, looping or not and prominent; outer intramarginal vein 0.5–1.5 mm from leaf margin, as prominent as tertiary venation.
Inflorescences terminal, dense metabotryoids, 2.5 cm long, peduncles 1 cm long, axes flattened, with 2 or 4 narrow wings, drying brown. Bracts c. 1.5 mm long, ovate, keeled, caducous; bracteoles 2 per flower, 1 mm long, similar to bracts.
Flowers c. 15 per inflorescence, within the inflorescence in triads, 4-merous, anthopodium absent, only known before anthesis, mature buds 5 × 3 mm. Hypanthium c. 4 × 3 mm, obconical, drying dark reddish brown, densely glandular-warty, hypanthium rim 2 mm long, glandular inside. Calyx lobes c. 1 × 2 mm, broadly rounded. Petals c. 3 × 3 mm, cucullate in bud. Stamens c. 100, filaments 2–3 mm long, anthers c. 0.4 mm long, ellipsoid. Ovary bilocular, surrounded by spongy tissue, ovules numerous per locule, ascending. Style 3–4 mm long, pointed.
Fruits 1-seeded, c. 20 × 25 mm, irregularly depressed globose, laterally compressed, green, drying black and, smooth, pericarp ± woody, 1 mm thick, hypanthium rim 1–2 mm long, 5–9 mm in diameter.
Seeds c. 15 × 20 mm, transverse ellipsoid, testa adhering to the pericarp, spongy inside and adhering to the outer surface of the cotyledons, cotyledons ± half-globose, facing surfaces undulate, outer surfaces densely echinate, protuberances obscured by spongy testa tissue.
The species is named after Eduard Ferdinand de Vogel (*1942). Ed de Vogel is a renowned authority on Malesian orchids, especially those from New Guinea. His contributions to the flora of Sulawesi are perhaps less well known: with almost 2000 specimens of excellent quality collected there in 1973–74 and 1979 – among them the type specimen of this species – he was one of the most prolific plant collectors on the island during the 20th century.
Flowering was recorded in August, fruiting in May.
Syzygium devogelii is endemic to the province of Central Sulawesi, currently known to occur in lower montane forest at two localities in LLNP from 1350–1400 m elevation (Figure
Syzygium devogelii has a limited geographical distribution and seems to be restricted to lower montane forest within a narrow elevational belt. Known from only two localities, the EOO and AOO cannot be estimated reliably for the species. Because of the low collection density in Central Sulawesi, we believe that the species is more widespread and common than it currently appears. Deforestation has been recorded close to the type locality (using the Global Forest Change website,
Most species of Syzygium are reported to have cotyledons with rather smooth outer surfaces, unlike the peculiar echinate cotyledons of S. devogelii. We here interpret the tissue covering the outer surface of the cotyledons (Figure
Juvenile specimens of Syzygium balgooyi are similar to S. devogelii in their leaf shape, colour, and venation. In fact, both species were treated as one morphotype in
(Paratypes). INDONESIA. Central Sulawesi (Sulawesi Tengah), LLNP, Kab. Poso, Kec. Lore Tengah: 9 km NW of Bariri, 100 m east of climate tower, tree-inventory plot Bariri NE, 1°39.4'S, 120°10.5'E, 1400 m, 9 Sep 2006: Culmsee H 1333 (sterile; BO [BO-1938455]!, CEB) & 1378 (sterile; CEB, K [K000993491]!); ibid. loco, 18 Aug 2011: Brambach F, Mangopo H, Firdaus, Faber M, Tiranda R 0818 (sterile; BO [BO-1938442]!, CEB, GOET [GOET020015]!) & 0845 (sterile; BO [BO-1938443]!, CEB, L!).
9 km NW of Bariri, 80 m south of climate tower, tree-inventory plot Bariri S, 1°39.5'S, 120°10.4'E, 1400 m, Jul 2006: Culmsee H 1252 (sterile; CEB, GOET [GOET020016]!) & 1564 (flower buds; CEB, L!).
Distribution map of four species of Syzygium in Central Sulawesi: Syzygium contiguum (orange diamonds), S. devogelii (yellow dots), S. eymae (light blue square), and S. galanthum (green triangles). Lore Lindu National Park is indicated by a black line. Map created with QGIS (
Syzygium eymae is characterised by small (usually 2.3–4 × 1.5–2.5 cm), (sub-)sessile, leaves with thickly coriaceous, (broadly) elliptic or obovate blades, dense terminal inflorescences, and small, pyriform flowers with a calyptrate calyx that bears a minute apical opening and splits irregularly at anthesis. It differs from the morphologically similar Syzygium paradoxum (Merr.)
INDONESIA. Central Sulawesi (Sulawesi Tengah), Kab. Tojo Una-Una, Border of Kec. Ulubongka and Kec. Ampana Tete, Mt Lumut, between summit and western secondary peak, c. 1°12.3'S, 121°47.6'E, ± 2200 m (“Selebes, Res. Menado. O.afd. Poso. G. Lóemoet, Pilaartop en W. bijtop. (summit)”), 5 Sep 1938: Eyma 3624 (flowers; holotype U [U.1439024]!; isotypes: BO [BO-1679767]!, L [L.2535689]!).
Trees, height, bark and wood unknown. Young branchlets slender, 0.5–1 × 1–2 mm, rectangular in cross section, ridges arising at the petioles and running downwards to next node, epidermis drying reddish black; remaining angular or becoming ± terete, bark reddish brown and scaly; with 1–2 pairs of minute cataphylls near the base of the current flush.
Leaves opposite, (sub-)sessile. Petioles 0.5–2 × 1–1.5 mm, absent or very short and stout, drying black. Blades (1.8–) 2.3–4 (–5.2) × (1.2–) 1.5–2.5 (–3.1) cm, ratio 1.3–2, (broadly) elliptic or (broadly) obovate, base obtuse or rounded, apex rounded, acute, or shortly acuminate, margin revolute; thickly coriaceous (c. 0.3 mm thick), dull, drying dark reddish grey to reddish black above, (very) dusky red beneath; without black gland dots. Midrib impressed above, prominent, rounded, and darker than the lamina beneath. Secondary vein pairs (4–) 5–7, 3–10 mm apart, channelled and inconspicuous above, (slightly) prominent and more reddish than the lamina beneath; intersecondary veins sometimes present. Tertiary veins reticulate, channelled above, indistinct beneath. Inner intramarginal vein 1–2 mm from leaf margin, looping; outer intramarginal vein not present.
Inflorescences terminal, 2-nodate metabotryoids, ≤ 3 cm long, peduncles ≤ 1 cm long, axes angular. Bracts c. 1–1.5 mm long, deltate, keeled, caducous; bracteoles 2 per flower, c. 1 mm long, similar to bracts.
Flowers ≤ 10 per inflorescence, within the inflorescence in triads, anthopodium absent, mature buds 5–6 × 3 mm. Hypanthium 4–5 × 4–5 mm, pyriform, smooth, hypanthium rim 1.5 mm long. Calyx lobes calyptrate with small apical opening, slightly lighter-coloured than hypanthium when dry, splitting irregularly at anthesis, caducous. Petals calyptrate, adhering to the calyx. Stamens c. 50, filaments 6–7 mm long, white, anthers c. 0.4 mm long, ellipsoid. Ovary bilocular, ovules several per locule, ascending. Style 6–7 mm long, pointed.
Fruits and seeds unknown.
The species is named after Pierre Joseph Eyma (1903-1945), one of the early botanists to explore the mountainous regions of Central Sulawesi (
The species was collected in flowering state in September 1938.
S. eymae is endemic to the province of Central Sulawesi and currently only known from the type locality: Mt Lumut on Sulawesi`s eastern peninsula (Figure
With only the type specimen known, we consider S. eymae “Data Deficient” (DD) at present, following the IUCN Red List Categories and Criteria (
The species of tribe Syzygieae Wilson (in
The flowers of Cleistocalyx are described as having “calyptrate calyces, the undivided, often more or less indurated upper parts of which fall as a lid”, the lid often remaining attached at one side of the flower at early anthesis (
Most species of Syzygium with calyptrate calyces are clearly different from S. eymae in their much larger leaves with more pairs of secondary veins. The few small-leaved species can all be easily distinguished: S. paradoxum from Borneo differs by the characters given in the diagnosis. S. pseudocalcicola Craven & Biffin (in
Syzygium eymae is also superficially similar S. paucivenium (Merr.)
Several specimens collected on Mt Rorekautimbu in LLNP at 2400 m (e.g. Brambach et al. 0768) may belong here. They are morphologically similar to the type specimen, but have longer petioles. Since we currently lack flowering material of these specimens and because of the large distance between the respective collection localities, we prefer to await more specimens before incorporating these collections in S. eymae.
“Myrtaceae sp. 7” (
Syzygium galanthum is similar to Syzygium hylochare (Diels)
INDONESIA. Central Sulawesi (Sulawesi Tengah): LLNP, Kab. Poso, Kec. Lore Tengah, 3.5 km NE of Rompo, following road to Katu for 3 km, then following footpath N for 2 km, tree-inventory plot Tarara, 1°35.3'S 120°17.0'E, 1200 m, 29 Nov 2011: Brambach F, Mangopo H, Firdaus, Faber M, Tiranda R 1316 (flowers; holotype L[L.3962132]!; isotypes BO [BO-1938381]!, CEB, GOET [GOET020017]!, K [K000993484]!).
Trees, up to 25 m tall, diameter at breast height ≤ 30 cm, trunk straight, ≤ 15 m tall, with buttresses 0.4 m tall, sometimes with stilt roots. Outer bark bright- or rusty red, peeling off in thin sheets, inner bark pale or dark red, wood straw or cream-coloured. Young branchlets 1–2 × 2–3 mm, subangular, flattened, epidermis olive, drying reddish brown, striate; becoming ± terete, bark (reddish) brown, striate or fissured, later peeling off in small thin sheets.
Leaves (sub-)opposite. Petioles 6–18 × 1–3 mm, channelled above, rounded beneath, turning corky, pale brown, drying (reddish) brown. Blades (10–) 12–23 (–26) × (4–) 5.5–7.5 (–9) cm, ratio (1.7–) 2.5–3.2 (–3.5), (narrowly) elliptic or rarely oblanceolate, base acute, obtuse, or rounded, apex acuminate, acumen often recurved, margin flat or revolute; chartaceous, glossy green and often ± bullate above, paler green beneath, drying dull and (greyish or olive) brown above, dull and (yellowish or greyish) brown beneath; pellucid dots scattered or numerous, usually visible on lower surface and sometimes also on upper surface. Midrib channelled above, prominent, rounded, drying pale or reddish brown, striate and with dark gland dots beneath. Secondary vein pairs (6–) 8–10 (–12), 7–25 mm apart, prominulous or not, concolorous with the lamina and usually inconspicuous above, prominent and concolorous with or more reddish than the lamina beneath; intersecondary veins present. Tertiary veins reticulate, lax, prominulous or not, concolorous with the lamina and usually inconspicuous above, prominent and concolorous with or more reddish than the lamina beneath. Inner intramarginal vein 3–8 mm from leaf margin, (strongly) looping; outer intramarginal vein 1–3 mm from leaf margin.
Inflorescences axillary on leafless portion of the twigs, often fascicled, lax, (sub-)sessile botryoids or monads, 3–5 cm long, peduncles absent or ≤ 1 cm long, axes angular, drying (reddish) brown with many black gland dots, turning corky at the base, often with conspicuous whitish blisters. Bracts c. 0.5 mm long, early caducous; bracteoles 2 per flower, similar to bracts.
Flowers 1–8 per inflorescence, within the inflorescence in monads, 4-merous, only known before anthesis, mature buds 15–25 × 5–7 mm, anthopodium 5–10 (–14) mm long, slender. Hypanthium 7–11 × 5–7 mm, infundibuliform, pale green, drying dark reddish brown, wrinkled, densely black gland-dotted and with conspicuous whitish blisters, hypanthium rim 3 mm long. Calyx lobes 2 × 3–5 (outer) and 3–4 × 5–7 (inner) mm, broadly rounded with thin hyaline margins, greenish white, drying red, sparsely gland-dotted. Petals c. 8 × 6 mm, hood shaped before anthesis, milky white, drying yellowish red, faintly veined and densely pellucid-dotted. Stamens c. 100, filaments 4–10 mm long before anthesis, yellowish green, anthers c. 1 mm long, ovoid or ellipsoid, yellow. Ovary bilocular, locules surrounded by spongy tissue, ovules many per locule, ascending. Style 10 mm long before anthesis, pointed, green.
Fruits and seeds unknown.
The species name derives from the Greek γάλα (milk) and άνθος (flower) and refers to the petals’ milky white colour (Figure
The type specimen was collected with mature flower buds in late November, suggesting flowering in December.
Syzygium galanthum is currently only recorded from LLNP in the province of Central Sulawesi (Figure
Syzygium galanthum has a limited geographical distribution (estimated EOO 140 km²) and seems to be restricted to submontane forest between 700 and 1200 m. We assume that the estimated AOO of 12 km² is unrealistically low, due to limited collection activities in Central Sulawesi. However, despite being inside the protected LLNP, recent deforestation activities have been detected near one of the collection sites (Pono inventory plot, detected using the Global Forest Change website,
Tambeanitu (Bahasa Behoa, Brambach et al. 1316).
In the field, S. galanthum can be recognised by the leaves with corky petioles and rather few, ± arching secondary veins. Similar corky petioles occur in S. peregrinum (Blume) Merrill & Perry (1939, 154) from Borneo and the Southern Philippines. A peculiarity is the presence of white blisters on the inflorescence axes and flowers of dried material (Figure
It appears that there is a group of morphologically similar species in Malesia, all characterised by pale-drying leaves with rather few secondary veins, inflorescences below the leaves, and medium-sized to large, showy, infundibuliform flowers with short or long anthopodia and either white or red/pink petals and stamens: e.g. S. iliasii
(Paratypes). INDONESIA. Central Sulawesi (Sulawesi Tengah), LLNP: Kab. Poso, Kec. Lore Tengah, 3.5 km NE of Rompo, following road to Katu for 3 km, then following footpath N for 2 km, tree-inventory plot Tarara, 1°35.3'S 120°17.0'E, 1200 m, 22 Nov 2011: Brambach F, Mangopo H, Firdaus, Faber M, Tiranda R 1047 (sterile; BO [BO-1938446]!, CEB, GOET [GOET020018]!) & 1083 (sterile; BO [BO-1938445]!,CEB, L!) & 1290 (sterile; BO [BO-1938444]!, CEB, K [K000993485]!).
Kab. Sigi, Kec. Kulawi, 2.4 km ENE of Toro, NE edge of Pono Valley, tree-inventory plot Pono, 1°29.7'S, 120°03.4'E, 1050 m, 16 Aug 2006: Culmsee 537 (sterile; CEB, K [K000993492]!) & 890 (sterile; BO [BO-1938448]!, CEB); ibid. loco, Jul 2007: Culmsee r497 (sterile; BO [BO-1938449]!, CEB, GOET [GOET020019]! , L!).
Kab. Sigi, Kec. Kulawi Selatan, 4 km ENE of Watukilo, 400 m N of Mboe River, tree-inventory plot Rantena, 1°36.2'S, 120°04.5'E, 700 m, 17–26 Jun 2011: Brambach F, Mangopo H, Firdaus, Faber M, Tiranda R 0533 (sterile; BO [BO-1938447]!, CEB, L!).
Morphological characters of Syzygium galanthum. a trunk, c. 15 m tall b upper side of leaves c branchlet tip with smooth younger petioles d older corky petioles e underside of leaf f bark slash g mature flower buds in fresh state h dried mature flower bud with apical part of inflorescence axis and white blister on the anthopodium i longitudinal section of mature flower bud in dried state j branch with mature flower buds below the leaves k detail of dried, fascicled inflorescences. a–b and f–k type collection Brambach et al. 1316c and eBrambach et al. 1083dBrambach et al. 1047. All scale bars: 1 cm.
Leaves of all new species described. Variation of Syzygium balgooyi (a–e), S. contiguum (f), S. devogelii (g), S. eymae (h), and S. galanthum (i). aBrambach et al. 0283bBrambach et al. 0681cBrambach et al. 1333dde Vogel 5413 [L.2517563] eCulmsee r2162fCulmsee r463gBrambach et al. 0818hEyma 3624 [L.2535689] iBrambach et al. 0533. Scale bar: 10 cm, valid for all leaves.
We are indebted to Aiyen Tjoa, Hardianto Mangopo, Firdaus Dg. Matta, Rickson Tiranda, Dewi Ramadhaani, Ilfianti Kasmudin, and Matthias Faber for their valuable help during fieldwork. Sadie Barber (E) gave valuable advice for the photographs of dried specimens. We thank the LLNP authorities for allowing us access and RISTEK for their assistance with research and collection permits. The curators of the herbaria A, B, BM, BO, E, HBG, K, KY, L, M, U and US allowed us access to their collections, provided loans, or helped searching for specimens, for which we are grateful. We also thank Peter G Wilson for reviewing the manuscript and for his helpful suggestions to improve it. Fieldwork in Indonesia and herbarium work in the Netherlands and UK was funded by the German Research Foundation (DFG), grant CU127/3-1, and FB was supported by a PhD scholarship of Evangelisches Studienwerk Villigst (Germany). Both are gratefully acknowledged, as well as support by the Open Access Publication Funds of the Göttingen University.