Fernandoa adenophylla (Wall. ex G. Don) Steenis, Blumea 23(1): 135. 1976; et Fl. Males., Ser. 1, Spermat. 8(2): 158. 1977; Kochummen in Ng, Tree Fl. Malaya 3: 39. 1978; Santisuk & J. E. Vidal in J.-F. Leroy, Fl. Cambodge Laos Vietnam 22: 39. t. 5, fig. 1. 1985; Santisuk in Smitinand & K. Larsen, Fl. Thailand 5(1): 47. 1987.

Ngernsaengsaruay, Chatchai, Meeprom, Nattanon, Boonthasak, Weereesa, Attasook, Yanatshara & Thunthawanich, Raweewan, 2023, A taxonomic revision of Thai Fernandoa Welw. ex Seem. (Bignoniaceae), PhytoKeys 235, pp. 249-270 : 249

1. Fernandoa adenophylla (Wall. ex G. Don) Steenis, Blumea 23(1): 135. 1976; et Fl. Males., Ser. 1, Spermat. 8(2): 158. 1977; Kochummen in Ng, Tree Fl. Malaya 3: 39. 1978; Santisuk & J. E. Vidal in J.-F. Leroy, Fl. Cambodge Laos Vietnam 22: 39. t. 5, fig. 1. 1985; Santisuk in Smitinand & K. Larsen, Fl. Thailand 5(1): 47. 1987.

Figs 1 , 2

Bignonia adenophylla ≡ Bignonia adenophylla Wall. [Numer. List: 221. Wallich Cat. 6502, nom. nud.] ex G. Don, A Gen. Hist. 4: 221. 1838.

Spathodea adenophylla ≡ Spathodea adenophylla (Wall. ex G. Don) DC., Prodr. 9: 206. 1845.

Heterophragma adenophyllum ≡ Heterophragma adenophyllum (Wall. ex G. Don) Seem., J. Bot. 8: 340. 1870.

Haplophragma adenophyllum ≡ Haplophragma adenophyllum (Wall. ex G. Don) Dop, Bull. Soc. Bot. France 72: 890. 1925.

Type.

Myanmar, Ava , 12 Oct 1826, Wallich 6502A (lectotype, first step designated by van Steenis (1976: 135), G [without barcode], second step designated here G [G00133642, photo seen]; isolectotypes K-W [K001124064, photo seen], G [G00134691, G00134695, photos seen], P [P00609736, photo seen].

Description.

Trees deciduous, 5-15(-20) m tall, up to 170 cm girth; bark irregularly cracked, corky, grey to greyish-brown; young branches densely stellate and dendroid tomentose. Leaves decussate; petioles very short or up to 1.5 cm long; rachises 6-41 cm long, 4-angular, channelled above; petioles and rachises densely stellate and dendroid tomentose; leaflets 5-9, opposite, laminas variable in shape, size, apex and base, obovate, elliptic, ovate, oblong, suborbicular or orbicular, the terminal leaflets largest, 10.5-46.5 × 6.5-28 cm, the lateral leaflets 3-33 × 3-21 cm, the lowest pair of leaflets near the base of petiole much reduced, resembling foliaceous pseudostipules, 0.5-8.5 × 0.5-7 cm, apex obtuse, rounded, acute, acuminate, cuspidate or emarginate, base cuneate, oblique, obtuse, truncate, cordate or subcordate, margin entire or repand, subcoriaceous, glabrous above, except densely stellate and dendroid tomentose along midrib and basal part of lateral veins above, densely stellate and dendroid tomentose below, with scattered glands on both surfaces, midrib and lateral veins raised below, lateral veins 4-11 pairs, curving and connected in loops near the margin, veinlets reticulate, with small hairy domatia in the axil of lateral veins below; petiolules very short. Inflorescence a thyrse, 10-65 cm, erect, lax-flowered; peduncles 2-10 cm long; rachises 10-53 cm long; peduncles, axes and pedicels with dense stellate and dendroid tomentose. Flowers: calyx yellowish-green, thick, 5-ribbed, persistent, densely yellowish-brown dendroid tomentose outside, glabrous inside, campanulate, 2-4.5 × 1.5-3 cm, bilabiate, 5-lobed, upper lobes (posterior) 3 and lower lobes (anterior) 2, subequal or unequal, lobes triangular, 0.7-1.8 × 0.4-1.7 cm, apex acute; corolla yellowish-green, creamy white to pale yellow, thick, densely yellowish-brown dendroid tomentose outside, glabrous inside, corolla tube curved, constricted between basal and upper parts, basal tube short cylindrical, widened towards the base, 1.5-2.5 cm long, 1-2.5 cm wide at base, upper tube widened towards the mouth, campanulate, 3-5.5 cm long, 3.5-5 cm wide at mouth, bilabiate, 5-lobed, upper lobes 2 and lower lobes 3, subequal or unequal, lobes suborbicular or broadly obovate, 2.5-4 × 2.4-5.5 cm, apex rounded, crisped; stamens 4, didynamous, subexserted, longer pair 3.2-5.5 cm long, shorter pair 3-4.5 cm long, filaments arcuate, creamy white to pale yellow, glabrous, anthers 5-9 mm long; staminode 1, needle-like, 0.7-2.2 cm long; disc annular, surrounding the base of ovary, creamy white to pale yellow; ovary cylindrical, 0.6-1.5 cm long, with (6-)10 longitudinal ridges, densely dendroid tomentose, style slender, 3-5 cm long, creamy white to pale yellow, glabrous, stigma 2-lobed, 4-7 mm long. Fruits cylindrical, 34-85 × 1.5-3.5 cm, green turning brown when dry, twisted, with (6-)10 prominent longitudinal ridges, densely yellowish-brown dendroid tomentose, septum 2-3 mm thick, 1-1.5 cm wide. Seeds flat, rather rectangular with a lateral hyaline-membranous wing, 1.5-4 × 0.6-1.4 cm.

Distribution.

India (Assam, Andaman and Nicobar Islands), Pakistan, Bangladesh, Myanmar, Vietnam, Laos, Cambodia, Thailand, Peninsular Malaysia.

Distribution in Thailand.

NORTHERN: Mae Hong Son, Chiang Mai, Chiang Rai, Nan, Lamphun, Lampang, Tak, Sukhothai, Phitsanulok, Kamphaeng Phet, Nakhon Sawan; NORTH-EASTERN: Phetchabun, Loei, Sakon Nakhon, Khon Kaen; EASTERN: Nakhon Ratchasima, Ubon Ratchathani; SOUTH-WESTERN: Uthai Thani, Kanchanaburi, Phetchaburi, Prachuap Khiri Khan; CENTRAL: Suphan Buri, Saraburi, Bangkok (Queen Sirikit Park, cultivated); SOUTH-EASTERN: Chon Buri, Rayong, Chanthaburi, Trat; PENINSULAR: Chumphon, Ranong, Surat Thani, Phangnga, Nakhon Si Thammarat, Trang. (Fig. 3 )

Habitat and ecology.

It is found in deciduous dipterocarp and mixed deciduous forests (with or without bamboo), mixed deciduous forests with bamboo on limestone hills, savannas, gaps or edge of dry evergreen and lower montane rain forests, transition between deciduous and evergreen forests, secondary forests, disturbed open areas, along roadsides, along riverbanks, at elevations of near above mean sea level (a.m.s.l.) up to 1,000 m.

Phenology.

Flowering and fruiting nearly all year round.

Conservation status.

Fernandoa adenophylla is widely distributed from India to Indochina and Peninsular Malaysia, and has a large extent of occurrence (EOO of 3,353,755.49 km2) and area of occupancy (AOO of 352 km2). Also, considering it grows in secondary forests, disturbed open areas, and along roadsides, it is considered here as Least Concern (LC).

Etymology.

The specific epithet of Fernandoa adenophylla is derived from the Greek compound words, "aden-", "adeno-" meaning gland (glandular-), and "- phylla " meaning -leaved, refers to the leaves of this species with a few scattered glands on both surfaces ( Stearn 1992; Radcliffe-Smith 1998; Gledhill 2002).

Vernacular name.

Khae khon (แคขน) (Northern); Khae bit (แคบิด) (Northern, North-Eastern); Khae phong (แคพอง) (Peninsular, Surat Thani); Khae rao (แคร้าว), Khae lao (แคลาว) (North-Eastern, Loei); Khae hua mu (แคหัวหมู) (Nakhon Ratchasima); Khae hang khang (แคหางค่าง) (General); Haeng pa (แฮงป่า) (Chanthaburi); Hong pa (โฮงป่า) (South-Western); Karen wood, Katsagon, Katsagon tree, Petthan (common name); Dhopa-paroli (Assam); Marodphali (Hindi).

Uses.

Flowers and young fruits are consumed as boiled or grilled vegetables and required cooking. Cultivated as shade and ornamental trees (the author’s observations). The wood is locally used in construction and used for making farming utensils. Bark, leaves, and seeds are used as medicinal purposes ( Widodo 1998; Biodiversity-Based Economy Development Office (Public Organization) [BEDO] 2021).

Notes.

Bignonia adenophylla was named by Wallich based on Wallich Cat. 6502 collected from Myanmar: 6502A from Irrawaddy River, Yenangheum (Yenangyaung), Prome, Sagaen (Sagaing), and Ava and 6502B from Taong Dong but unpublished, and then this name was described by Don (1838: 221). van Steenis (1976: 135) mentioned Wallich Cat. 6502 of Ava and Prome as type, when he transferred B. adenophylla under Fernandoa adenophylla . Wallich Cat. 6502 represents two gatherings (two different materials collected from two different cites, which are distinguished by Wallich Cat. 6502A and Wallich Cat. 6502B, respectively). Wallich Cat. 6502A is from Ava and Prome and Wallich Cat. 6502B is from Taong Dong. Thus, Wallich Cat. 6502A could be regarded as the true type specimen, and Wallich Cat. 6502B from Taong Dong is not type.

van Steenis (1976) cited Wallich Cat. 6502 from Ava and Prome at G as the holotype with isotypes in K and P. However, Wallich’s collection numbers are known to be curated by species generally from multiple collections ( Noltie and Watson 2021); therefore, his erroneous designation of holotype effectively selected a) a collection for the lectotype if the collection consists of multiple sheets or b) the lectotype if it is a unicate gathering. van Steenis (1976) did not mention the number of specimens, and following Art. 9.6 of the ICN ( Turland et al. 2018), they constitute syntypes. Therefore, the name Fernandoa adenophylla has been lectotypified in a first step by van Steenis (1976) using specimen Wallich Cat. 6502A at G [without barcode] with isolectotypes at K [without barcode] and P [without barcode]. We located three sheets of the specimen Wallich 6502A from Ava at G [G00133642, G00134691, and G00134695] and two sheets of the specimen Wallich 6502A from Prome at G [G00133632 and G00134708]; The G [G00133642] specimen is better preserved and more complete than the others, and hence is selected here in a second step lectotypification. We also traced isolectotypes at K-W [K001124064] and P [P00609736].

Santisuk (1987) reported that the leaves, calyx, and ovary have a stellate tomentum, but in this study the calyx, corolla, ovary, and fruits only exhibited dendroid trichomes, whereas the leaves bear both stellate and dendroid trichomes.

In addition to Santisuk (1987), the distribution in Mae Hong Son, Chiang Rai, Phayao, Nan, Lamphun, Sukhothai, Phitsanulok, Kamphaeng Phet, Nakhon Sawan, Sakon Nakhon, Khon Kaen, Ubon Ratchathani, Uthai Thani, Suphan Buri, Saraburi, Bangkok (cultivated), Rayong, Ranong, and Nakhon Si Thammarat Provinces were newly recorded in this study.

Santisuk (1987) reported that this species occurs in deciduous forests, savannas, and lowland secondary forests. Further, we confirmed it grows in more variable habitats (see habitat).

Additional specimens examined.

Thailand. Northern: Mae Hong Son [Sop Moei, 12 Jul 2013, Pongamornkul 3468 (QBG); Sop Moei, 1 Jan 2014, Pongamornkul 3928 (QBG); Ban Mueang Paeng, Mueang Paeng Subdistrict, Pai District; Salawin Forest Plantation, Mae Sariang District (Ngernsaengsaruay own observation)]; Chiang Mai [Locality not specified, 4 Oct 1921, Nai Noi Mao s.n. (BK) ( Santisuk 1974); Chiang Dao District, fr., 27 Nov 1963, Bunchuai 1375 (BKF); Mae Rim District, Mae Sa valley, in cultivation area, 1,000 m alt., fl., 15 May 1974, Jackson 6039 (BKF); Hang Dong District, along the dirt road from Mae Heeyah Nai to Ban Bong, south side of Doi Suthep-Pui National Park, 425 m alt., fl., 15 Jul 1989, Maxwell 89-903 (L [L3730390]); Doi Chiang Dao, SE foothills near Ban Yang Pong Luang, Chiang Dao District, 525 m alt., fl., 29 Oct 1989, Maxwell 89-1326 (L [L3730538]); Doi Suthep-Pui National Park, Mae Rim District, in open place near Mae Sa stream, 650 m alt., fr., 9 Jan 1990, Maxwell 90-37 (L [L3730536, L3730537]); Department of Biology, Faculty of Science, Chiang Mai University, 350 m alt., fl., 27 Jun 1992, Palee 45 (L [L3730324]); Queen Sirikit Botanic Garden, Mae Rim District, 10 Aug 1994, BGO. Staff 1294 (QBG); Maesa Elephant Camp, Mae Rim District, 25 Aug 1994, BGO. Staff 1454 (QBG); Huai Gayo village area, San Kamphaeng District, 675 m alt., fr., 10 Oct 1995, Palee 325 (BKF, L [L3730483, L3730484]); Mae Rim District, 5 Nov 1997, Watthana & Siriphum 29 (QBG); Doi Suthep, Kerr 3262 (ABD, BM, K) ( Santisuk 1974)]; Chiang Rai [Doi Tung, base of the east side at Bah Kah (Akha) village, Mae Fa Luang District, 500 m alt., fl., 21 Jul 2006, Maxwell 06-494 (L [L3730659]); Mueang Chiang Rai District (Ngernsaengsaruay own observation)]; Phayao [Rom Yen Subsistrict, Chiang Kham District, 18 Jun 2013, La-ongsri et al. 2920 (QBG)]; Nan [Mueang Nan District, 14 Nov 2009, Srithi 345 (QBG); Song Khwae District, 18 Oct 2019 (the authors own observation)]; Lamphun [Mae Tha District, 29 Jan 2010., Romkham 132 (QBG)]; Lampang [Ngao, Mae Huat, 350 m alt., fl., 18 May 1954, Smitinand 1579 (BKF)]; Tak [along Mae Ping River, Om Lu Rapids, fl., fr., 9 Dec 1920, Rock 629 (US [US03206561, US03206562, US03206563]); Doi Pha Wo, trail from Nam Dip to Pang Luang, Mae Sot District, fl., 19 Dec 1920, Rock 669 (US [US03206564]); between Palut and Nam Dip, on the trail from Raheng to Mae Sot, fl., 17 Dec 1920, Rock 1080 (US [US03206556]); Lan Sang National Park, bamboo rich gallery forest along cascade, fr., 27 Dec 1974, Geesink, Hiepko & Phengklai 7941 (L [L2815249]); Lan Sang National Park, 300 m alt., fl., 5 Aug 1997, Setbubpa 19 (BKF); Ban Tak District, 16 Oct 2014, Tanming 715 (QBG); Mae Kor, Winit 410 (ABD, K) ( Santisuk 1974)]; Sukhothai [Si Satchanalai National Park, Si Satchanalai District, fl., 27 Jul 2015, Maknoi 7946 (BKF, QBG)]; Phitsanulok [Thung Salaeng Luang National Park, c. 80 km east of Phitsanulok, c. 500 m alt., fl., 25 Jul 1973, Murata, Fukuoka & Phengklai T-17126 (BKF, L [L2815258])]; Kamphaeng Phet [Mae Wong National Park, Khlong Lan District, 20 Oct 2001, Watthana 1528 (QBG); Wang Khuang Subdistrict, Phran Kratai District (Ngernsaengsaruay own observation)]; Nakhon Sawan [Takhli District, 30 Jan 2013, Maknoi 5210 (QBG)]; North-Eastern: Phetchabun [Khao Hin Pakarang Chon Daen District, 24 Nov 2009, Maknoi & Tanaros 3425 (QBG); Chon Daen District, Vacharapong 320 (BK) ( Santisuk 1974)]; Loei [Wang Saphung District, 300 m alt., fl., 28 Apr 1946, Nakkarn 66 (BKF); Wang Saphung District, 300 m alt., fl., y. fr., 28 Nov 1957, Bunpheng 1030 (BKF, K) ( Santisuk 1974); Pak Chom District, fl., 2 Sep 1968, Phengnaren & Smitinand 583 (BKF); Pha Daeng, Wang Saphung District, Suvarnakoses 1327 (BKF, K) ( Santisuk 1974)]; Sakon Nakhon [Phu Phan National Park (Ngernsaengsaruay own observation)]; Khon Kaen [Ban Na Chan, Phu Pha Man National Park, 477.5 m alt., fr., 25 Feb 2011, Norsaengsri, Tathana & Lakert 7616 (BKF, QBG)]; Eastern: Nakhon Ratchasima [Wang Nam Khiao District, Sakaerat Environmental Research Station, 420 m alt., fl., 17 Jul 1967, Damrongsak 115 (BKF); Wang Nam Khiao District, fr., Dec 1967, Damrongsak 418 (BKF); Khao Phayom, Wang Nam Khiao District, 430 m alt., fr., 21 Dec 1967, Anan 15 (BKF); Pak Thong Chai District, Lam Phra Phloeng Dam, 400 m alt., fr., 29 Jan 1983, Koyama, Terao & Wongprasert T-33101 (BKF); Nong Ra Wiang Subdistrict, fl., 3 Oct 2000, Garcia & Phengkhlai 385 (BKF); Ban Badan, Pak Thong Chai District, Sono 8 (BKF) ( Santisuk 1974); Khao Phayom, Wang Nam Khiao District, Nalampoon 15 (BKF) ( Santisuk 1974)]; Ubon Ratchathani [Khong Chiam District (Ngernsaengsaruay own observation)]; South-Western: Uthai Thani [Khao Pla Ra, Lan Sak District, 19 Sep 2015, Tanming 892 (QBG); Mueang Ka Rung Subdistrict, Ban Rai District; Phai Khiao Subdistrict, Sawang Arom District (Ngernsaengsaruay own observation)]; Kanchanaburi [Khwae Noi River Basin, near Neeckey (N. Wangka), 150 m alt., fr., 1 May 1946, Bloembergen & Kostermans 276 (BK, K, L [L2815284, L2815285], P [P02902332]); Khwae Noi River Basin, Kin Sai Yok, c. 120 km northwest of Kanchanaburi, along river bank, 100-150 m alt., fl., 2 Aug 1946, Kostermans 1436 (L [L2815280, L2815281], US [US03206558]); Ban Kao, fl., 14 Nov 1961, Larsen 8202 (L [L2815289]); Sai Yok District, sterile, 15 Dec 1961, Larsen 8733 (L [L2815287, L2815288]); Sai Yok District, 150 m alt., fl., 23 Nov 1971, van Beusekom et al. 3907 (BKF, L [L2815283]); Huai Ban Kao, 750 m alt., fr., 9 Nov 1971, van Beusekom et al. 3576 (BKF, L [L2815277]); Forest Research and Demonstration Center, Chalae Subdistrict, Thong Pha Phum District, 700 m alt., fr., 11 Dec 1995, van Welzen 2 (L [L2815250]); Bo Phloi, Lam I Su, fr., 13 Apr 2000, Phengklai et al. 12171, 12172 (BKF); Wang Pho, Lum Sum Subdistrict, Sai Yok District, 50 m alt., fl., 1 Jul 2006, Chongko 532 (L [L3731040, L3731041], QBG); Mahidol University, Kanchanaburi Campus, Sai Yok District, 21 Jul 2006, Maxwell 06-494 (QBG); Thipsukhontharam Temple, Huai Krachao District, 14 Oct 2014, Tanming 633 (QBG); Khao Thong, Kerr 19627 (ABD, BK, BM) ( Santisuk 1974)]; Phetchaburi [Khao Phanoen Thung, Kaeng Krachan National Park, 23 Aug 1998, Sasirat 80 (QBG); Thung Luang, Kerr 20638 (ABD, BK, BM) ( Santisuk 1974)]; Prachuap Khiri Khan [Bang Saphan District, Put 1383 (BK, BM) ( Santisuk 1974)]; Central: Suphan Buri [Phu Hang Nak, U Thong District (Ngernsaengsaruay own observation)]; Saraburi [Sam Lan Forest, Mueang Saraburi District, 75 m alt., fl., 27 Jul 1975, Maxwell 75-722 (L[L2815259])]; Bangkok [Queen Sirikit Park, cultivated (Ngernsaengsaruay own observation)]; South-Eastern: Chon Buri [Si Racha District, sterile, 1 Dec 1927, Collins 1748 (BK, US [US03206557]); Khao Khiao, Si Racha District, 100 m alt., fl., 13 Jun 1976, Maxwell 76-393 (L [L2815252]); Khao Khiao Open Zoo, 9 Aug 2000, Phengklai 12610 (BKF); Si Racha District, Collins 223 (ABD) ( Santisuk 1974)]; Rayong [Klaeng District, 19 Dec 2007, Wessumritt 55 (QBG); Chanthaburi [Khao Sa Bap, Makham District, 11 May 1956, Chit 333 (BKF) ( Santisuk 1974)]; Trat [wayside near sea level, fl., 1 Aug 1973, Murata, Fukuoka & Phengklai T-17348 (BKF, L [L2815254, L2815255], Murata, Fukuoka & Phengklai T-17389 (BKF, L [L2815251])]; Peninsular: Chumphon [Tha Sae District, Jaray 110 (BK) ( Santisuk 1974); Kuring, Kerr 11609 (BK, K, L [L2815292]) ( Santisuk 1974)]; Ranong [30-70 km south of Ranong, 50-100 m alt., fl., 27 Apr 1974, Larsen & S. S. Larsen 33441 (BKF, L [L2815256])]; Surat Thani [Ban Na San District, 100 m alt., fl., 14 Oct 1957, Thaworn 500 (BKF, K) ( Santisuk 1974)]; Phangnga [Nang Yon, Takua Pa, District, Kerr 17047 (ABD, BK, BM) ( Santisuk 1974)]; Nakhon Si Thammarat [Khiri Wong, Lan Saka District, fl., 18 Feb 1962, Ploenchit 1792 (BKF); Tha Sala District, Walailak University, 10 m alt., fl., 22 Dec 2006, Pooma, Pattharahirantricin & Sirimongkol 6528 (BKF); Trang [Khao Chong, Na Yong District, 80 m alt., fl., 29 Sep 1965, Bunnab 6 (BKF, L [L2815293]); Khao Chong, Na Yong District, 84 m alt., fl., y. fr., Sep 1965, Bunnab 193 (BKF); Huai Yot, 1 May 1916. Vanpruk 852 (BKF)].

Figures

Figure 1. Fernandoa adenophylla A habitat and habit B bark C branch and leaves D branches, leaves and inflorescences E inflorescences and flowers F branches, leaves, inflorescences and fruits G winged seeds. Photos: Yanatshara Attasook (A, B) Chatchai Ngernsaengsaruay (C - G).

Figure 2. Lectotype of Fernandoa adenophylla, Wallich 6502 A (G [G 00133642]) from Ava, Myanmar. Photo: Conservatoire et Jardin botaniques de la Ville de Geneve, Geneve, Switzerland https: // www. ville-ge. ch / musinfo / bd / cjb / chg / adetail. php? id = 55790 & lang = en.

Figure 3. Distribution of Fernandoa in Thailand: F. adenophylla occurs in all floristic regions of Thailand and F. collignonii known only from Nan Province, Northern Thailand. [Thailand floristic regions follow Flora of Thailand Volume 16 Part 1 (The Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation 2022)].