Description
Elaeagnus ×maritima Koidz., Bot. Mag. (Tokyo) 31: 133, 1917.—TYPE: JAPAN. Kanagawa Pref., Sagami, Hayama, 3 Nov 1916,
K.Hisautsi s.n. (lectotype, TI, seen as photo!, designated here, see
Fig. 4A); Musashi, Yokohama, 5 Nov 1916,
K.Hisautsi s.n. (syntype, TI, seen as photo!).
Elaeagnus ×
hisauchii Makino ex Nakai, Bot. Mag. (Tokyo) 32: 223, 1918.—TYPE: KOREA. Jeju-do, 2 Nov 1917,
T.Nakai 6352 (syntype, TI, seen as photo!, see
Fig. 4B. JAPAN. Musashi, Yokohama, 5 Nov 1916,
K.Hisautsi s.n. (syntype, TI, not seen); Yokosuka,
J.Matsumura s.n. (syntype, TI, not seen).
Elaeagnus ×liukiuensis Rehder, J. Arnold Arbor. 1: 181, 1920.—TYPE: JAPAN. Okinawa (Liukiu), Okinawa Island, common round Naha, 1 Mar 1917, E.H.Wilson 8159 (holotype, A, seen as photo!; Isotype, K, US, seen as photo!).
Shrubs or vines, evergreen, about 3 m high. Stems scandent; bark shallowly or irregularly longitudinally fissured, gray or grayish brown; lenticels orbicular or elliptic, dark gray; branches gray or grayish brown, glabrous, without thorn; twigs prismatic, brown, grayish brown or gray, 1.5–2.3 mm wide, conspicuous ribs, densely reddish brown peltate scales; winter bud naked, terminal bud oblong or widely ellipsoid, 2.4–3.3 × 1.4–1.9 mm, lateral bud oblong or ovoid, 1.4–2.2 × 0.6–1.4 mm, brown to dark brown, densely reddish brown peltate scales. Leaves alternate, simple; petioles brown, grayish brown or gray, 11.3–19.2 × 1.2–2 mm, densely with reddish brown peltate scales; blades ovate-elliptic, broadly ovate, broadly elliptic or orbicular, 4.8–9.2 × 3.2–5.6 cm, apex acute to acuminate or often caudate with obtuse tip, margin entire, undulate, base obtuse to rounded, coriaceous, adaxial surface dark green, lustrous, sparsely brown peltate scales, glabrescent, abaxial surface yellowish green, densely reddish brown peltate scales or silvery white mixed with reddish brown peltate scales. Inflorescences axillary, fasciculate, 2–5 flowered; pedicels white or pale yellow, 2.6–4.6 × 0.5–0.7 mm, reddish brown peltate scales. Flowers bisexual, incomplete, apetalous, 3.9–6.2 mm in diameter; Calyx tubular, white or pale yellow, densely silvery white peltate scales mixed with reddish brown peltate scales; calyx tube rather broader, gradually constricted above ovary, 4–5.3 × 1.7–3.7 mm, 4-angled; 4-lobed, lobes triangular-ovate, 2– 3.6 × 1.7–3.5 mm, apex acute to acuminate, margin entire; stamens 4, epipetalous, antisepalous, borne in throat of calyx tube; filaments linear, white or yellowish white, ca. 0.4 mm long; anthers basifixed, narrowly oblong to elliptic, yellowish brown or brown, 1.9–2.2 × 0.8–1 mm; ovary epigynous, elliptic, 1.8–2.6 × 1–1.4 mm, 1-loculed; style linear, yellowish green or green, 4.3–5.1 mm long, glabrous or sparsely white stellate hair; stigma, clavate, bended, herkogamy absence or slightly approach hekogamy. Fruits drupe, ellipsoid, red-orange at maturity, 13.7–16.3 × 4.6–6.3 mm, silvery white peltate scales. Seeds oblong, yellowish brown, 13.3–16.1 × 4–5.8 mm, apex gradually constricted, obtuse, surface 8-grooved.
Korean name: Nok-bo-ri-ttong-na-mu (녹보리똥나무).
Phenology: Flowering October to November and fruiting April to May.
Distribution: Japan, Korea (Jeollanam-do, Gyeongsangnam-do, Jeju-do).
Habitat: Streamside, scrublands, and thickets.
Taxonomic notes: Elaeagnus ×
hisauchii was similar to
E. macrophylla by broadly ovate to nearly orbicular leaves and 4-angled campanulate calyx tube. However, it differed from
E. macrophylla by reddish brown twigs and petioles, short petioles, small leaves, and gradually constricted calyx tube above the ovary. Therefore, it was previously treated as a synonym of
E. ×
submacrophylla by having the abaxial surface of leaves with dense silvery white and sparse brown peltate scales, style with white stellate hairs (
Nakai, 1928;
Lee, 2003). However,
E. ×
hisauchii in Korea was observed not only on Jeju Island, which was previously known as type locality, but also on Heuksan-do Island (see
Fig. 4D), and these regions co-occur with
E. glabra and
E. macrophylla. Also, it differed from
E. ×
submacrophylla by having a vine and gradually constricted calyx tube above the ovary. Consequently,
E. ×
hisauchii in Korea should be considered a synonym of
E. ×maritima, which is regarded as a hybrid of
E. glabra and
E. macrophylla.
Elaeagnus ×
hisauchii was described by
Nakai (1918) based on a specimen from Jeju Island (
T.Nakai 6352, TI; Syntype, see
Fig. 4B). However, the specimen has been identified as
E. glabra. Rather, another specimen from Jeju Island (
T.Nakai 6351, TI, see
Fig. 4C) was identified as
E. ×
hisauchii and best represented the morphology mentioned in the original description. To accurately identify
E. ×
hisauchii, a lectotype among other syntypes should be designated. However, the lectotype could not be designated in this study because other syntypes could not be found.
Additionally,
E. ×maritima was assumed as a synonym of
E. glabra based on the morphological comparison (external morphology, trichome and pollen) (
Ki, 2004;
Koh, 2005), on the other hand, there was a view of integrating into
E. ×submacrophylla (
Lee, 1996). However, this species differs from
E. glabra by thick and angular twigs, more than 1 cm long petioles, ovate-elliptic, broadly elliptic, broadly ovate or orbicular leaves, and rather broader and 4-angled calyx tube. In addition, it was clearly distinguished from
E. ×
submacrophylla by the abaxial surface of the leaves with dense reddish brown scales or silvery white peltate scales mixed with reddish brown peltate scales, gradually constricted calyx tubes, and seeds without beaks (
Figs. 5,
6).
Representative specimens examined: KOREA. Jeollanam-do: Sinan-gun, Heuksan-myeon, Is. Heuksan, 31 Jan 2008, J.K. Ahn et al., SB20083 (KB); Sinan-gun, Heuksan-myeon, Ye-ri, 22 Oct 2011, B.U. Oh, Sinangun(Daeheuksando)-111022-095–096 (KH); Wando-gun, Bogil-myeon, Is. Bogildo, May 1965, L.M. Kim, KFR20015800–20015802 (KH); Wando-gun, Bogil-myeon, Yesong-ri, Suribong Peak, 7 Apr 2022, B.K. Park et al., Suribong-220407-027–028 (KH); Wando-gun, Gunoe-myeon, 25 Jun 2012, C.J. Oh et al., JN_V2012031 (KH); Wando-gun, Gunoe-myeon, Wando Arboretum, 4 Feb 2005, J.E. Koh & J.H. Kim, B200508052079, s.n. (SKK); Wando-gun, Gunoe-myeon, Daemun-ri, Wando Arboretum, 20 Jul 2013, M.S. Kang et al., 2013JNV027–028 (KH); Wando-gun, Gunoe-myeon, Yeongpung-ri, 3 Oct 2003, K.I. Heo et al. 27983, 28069, 28086, 28087, 28824 (SKK); Wando-gun, Gunoe-myeon, Yeongpung-ri, 23 Oct 2005, J.H. Lee et al. s.n. (SKK); Jindo-gun, Gogun-myeon, Jindo, 1969, L.M. Kim & M.Y. Cho, KFR20015666–20015668 (KH); Jindo-gun, Uisin-myeon, Temp. Ssanggyesa, 19 May 2002, K.I. Heo et al., 26687, 26690, 26691 (SKK); Jindo-gun, Uisinmyeon, Sacheon-ri, Mt. Cheomchalsan, Temp. Ssanggyesa, 6 Nov 2004, K.I. Heo & J.E. Koh s.n. (SKK); Jindo-gun, Uisin-myeon, Sacheon-ri, Unlimsanbang, 4 Nov 2004, K.I. Heo & J.E. Koh s.n. (SKK); Jindo-gun, Jindo-eup, Mt. Cheomchalsan, A path up a mountain of Ariranbi, 5 Nov 2004, K.I. Heo & J.E. Koh s.n. (SKK). Gyeongsangnam-do: Geoje-si, Nambu-myeon, Jeogu-ri, 22 Jun 2010, E.S. Jeon, ESJeon101305 (KH). Jeju-do: Mt. Hallasan, May 1966, L.M. Kim & S.G. Kang, KFR20015663–20015664, s.n. (KH); Jeju-si, Gujwa-eup, Songdang-ri, 21 Sep 2021, B.K. Park et al., Songdangri-210928-001 (KH); Jeju-si, Gujwaeup, Songdang-ri, 17 Oct 2021, K.H. Lee, Songdangri-211017-001–002 (KH); Jeju-si, Gujwa-eup, Songdang-ri, 23 Mar 2022, B.K. Park et al., Songdangri-220323-001–002 (KH); Gujwa-eup, Songdang-ri, 10 Oct 2022, E.S. Kang et al., Songdangri-221012-001–007 (KH); Jeju-si, Arang-dong, Yangji park, 7 Feb 2005, J.H. Kim et al. 054091, 054100, 054102, 054112 (SKK); Jeju-si, Hangyeongmyeon, Cheongsu-ri, Sanyang Keununggot, 2 Nov 2021, B.K. Park et al., Cheongsuri-211102-003–005 (KH); Jeju-si, Haean-dong (Gwangryeongcheon), 28 Sep 2012, D.S. Kim & S.Y. Kim, HALLA1701 (KH); Seogwipo-si, Sanghyo-dong, Donnaeko, 31 Oct 2005, J.H. Kim 04671, 04672, 054252 (SKK); Seogwipo-si, Sanghyo-dong, 23 Mar 2022, B.K. Park et al., Sanghyodong-220323-001–003 (KH); Seogwipo-si, Sanghyo-dong, 26 Apr 2022, Y.J. Jang & K.H. Lee, Sanghyodong-220407-007–008 (KH); Seogwipo-si, Jungmun-dong, Cheonjeyeon Waterfalls, 29 Oct 2010, C.H. Kim et al. 51828 (KB).
Elaeagnus ×submacrophylla Servett., Beih. Bot. Centralbl. 25: 84, 1909.—TYPE: JAPAN. Kyushu
, Nagasaki Pref., Nomo-saki,
Buerger 385 (lectotype, L, seen as photo!, designated here, see
Fig. 7A);
Buerger 384 (syntype, L, seen as photo!).
Elaeagnus ×nikaii Nakai, Bot. Mag. (Tokyo) 32: 224, 1918.—TYPE: JAPAN. Yamaguchi Pref., Nagato, Hagi, Tsuruedai, 3 Nov 1917,
J. Nikai 2723 (lectotype, TI, seen as photo!, see
Fig. 7B; isolecotype, TNS, seen as photo!).
Shrubs, evergreen, about 3 m high. Stems erect; bark shallowly or irregularly longitudinally fissured, gray or grayish brown; lenticels orbicular, dark gray; branches dark gray or grayish brown, glabrous, thorn present or absent; twigs terete, brown, grayish brown or gray, 1.6–2.2 mm wide, densely reddish brown peltate scales; winter bud naked, brown to dark brown, densely reddish brown peltate scales, terminal bud oblong or widely ellipsoid, lateral bud oblong or ovoid, 1.5–2.3 × 0.7–1 mm. Leaves alternate, simple; petioles brown, grayish brown or gray, 10.6–19.1 × 1.1–2.7 mm, densely reddish brown peltate scales; blades ovate-elliptic to broadly elliptic, 5.6–9.3 × 3.7–5.1 cm, apex acute to acuminate or often caudate with obtuse tip, margin entire, undulate, base obtuse to rounded, coriaceous, adaxial surface dark green, lustrous, sparsely silvery white peltate scales, glabrescent, reticulate vein inconspicuous when they are dry, abaxial surface yellowish green, densely silvery white peltate scales, punctured with reddish brown peltate scales. Inflorescences axillary, fasciculate, 1–3 flowered; pedicels white or pale yellow, 2.4–3.9 × 0.6–0.7 mm, reddish brown peltate scales. Flowers bisexual, incomplete, apetalous, 5.5–9.5 mm in diameter; Calyx campanulate, white or pale yellow, silvery white peltate scales and reddish brown peltate scales; calyx tube abruptly constricted above ovary, 4.9–6.2 × 2.7–4.2 mm, 4-angled; 4-lobed, lobes triangular-ovate, 2.7–4.7 × 2.2–4.4 mm, apex acute to acuminate, margin entire; stamens 4, epipetalous, antisepalous, borne in throat of calyx tube; filaments linear, white or yellowish white, 0.4–0.5 mm long; anthers basifixed, narrowly oblong to elliptic, yellowish brown or brown, 2.9–3.2 × 1–1.3 mm; ovary epigynous, elliptic, 2.2– 3.1 × 1.3–1.8 mm, 1-loculed; style linear, yellowish green or green, 8–9.3 mm long, whitish stellate hairs; stigma beyond stamens, clavate, bended, herkogamy absence or slightly approach hekogamy. Fruits drupe, ellipsoid, red-orange at maturity, 11.7–12.4 × 5.2–5.8 mm, silvery white peltate scales. Seeds oblong, yellowish brown, 10.7–11.2 × 4.2–4.8 mm, apex attenuate, beaked, surface 8-grooved.
Korean name: Keun-bo-ri-jang-na-mu (큰보리장나무).
Phenology: Flowering October to November and fruiting April to May.
Distribution: Japan, Korea (Gyeongsangnam-do).
Habitat: Streamside, scrublands, and thickets.
Taxonomic notes:
Elaeagnus ×submacrophylla was described by
Servettaz (1909) and is assumed to be a hybrid of
E. pungens and
E. macrophylla. However it was recognized as a hybrid of
E. glabra and
E. macrophylla by
Nakai (1928), who treated
E. ×
hisauchii Nakai as a synonym of
E. ×submacrophylla, and the hybrid of
E. pungens and
E. macrophylla was treated as
E. ×
nikaii Nakai (
Lee, 1996;
Lee, 2003;
Ohashi, 2003). In a recent taxonomic treatment,
E. ×
hisauchii was treated as a synonym of
E. ×maritima, which is regarded as a hybrid of
E. glabra and
E. macrophylla, and
E. ×
nikaii was treated as a synonym of
E. ×submacrophylla, which is regarded as a hybrid of
E. pungens and
E. macrophylla (
Ohashi, 2003;
Ko, 2015).
Elaeagnus ×submacrophylla in Japan has been known to be distributed in regions where
E. pungens and
E. macrophylla co-occur (
Ohashi and Yoshida, 2003). But
E. ×submacrophylla previously reported in Korea is confused about the distribution in that it was a region where
E. glabra and
E. macrophylla co-occur or only
E. macrophylla occurs (
Ko, 2015). As a result of the field survey and reclassification of the specimens, those identified as
E. ×submacrophylla in Korea were confirmed to be either
E. macrophylla or
E. ×
hisauchii (=
E. ×maritima). In addition,
E. ×
nikaii in Korea was not only previously reported on Mt. Mireuksan, Tongyeong where
E. pungens and
E. macrophylla co-occur, but also was morphologically similar to
E. macrophylla, but differed by reddish brown twigs, petioles, midrib of abaxial surface of leaves, style with no silvery white peltate scales and only white stellate hairs (
Figs. 8,
9). Therefore,
E. ×
nikaii previously reported in Korea is considered to be
E. ×submacrophylla.
Representative specimens examined: KOREA. Gyeongsangnam-do: Geoje-si, Dundeok-myeon, Sanbangri, Temp. Bohyeonsa, 30 Mar 2022, B.K. Park et al., Bohyeonsa-220330-001–002 (KH); Dundeok-myeon, Sanbang-ri, Temp. Bohyeonsa, 19 Oct 2022, Y.J. Jang et al., Bohyeonsa-221019-003–004 (KH); Tongyeong-si, Bongpyeong-dong, Mt. Mireuksan, 26 Apr 2005, J.H. Lee et al. 044754–044756, 046451–046453, 054328, s.n. (SKK); Tongyeong-si, Bongpyeong-dong, Mt. Mireuksan, 27 Oct 2021, B.K. Park et al., Mireuksan-211027-001–002 (KH); Tongyeong-si, Bongpyeong-dong, Mt. Mireuksan, 31 Mar 2022, B.K. Park et al., Mireuksan-220331-001–005 (KH); Tongyeong-si, Bongpyeong-dong, Mt. Mireuksan, 20 Oct 2022, B.K. Park et al., Mireuksan-220331-001–005 (KH); Tongyeong-si, Bongpyeong-dong, Mt. Mireuksan, 20 Oct 2022, Y.J. Jang et al., Mireuksan-220331-004 (KH).