Uvariodendron connivens (Benth.) R.E.Fr., Acta Horti Berg. 10: 55 (1930)

Dagallier, Leo-Paul M. J., Mbago, Frank M., Couderc, Marie, Gaudeul, Myriam, Grall, Aurelie, Loup, Caroline, Wieringa, Jan J., Sonke, Bonaventure & Couvreur, Thomas L. P., 2023, Phylogenomic inference of the African tribe Monodoreae (Annonaceae) and taxonomic revision of Dennettia, Uvariodendron and Uvariopsis, PhytoKeys 233, pp. 1-200 : 1

Uvariodendron connivens (Benth.) R.E.Fr., Acta Horti Berg. 10: 55 (1930)

Figs 7A-D , 15 , 16

≡ Uvaria connivens Benth., Trans. Linn. Soc. London 23(3): 465 (1862); Uva connivens Kuntze, Revis. Gen. Pl. 1: 8. (1891). Type. Equatorial Guinea - Bioko Sur • G. Mann 1159 (lectotype: K! (K000198803); isolectotypes: P! (P00362655), K! (K000198804, K000198805)), Fernando Po; 3°30'N, 8°40'E; 1861.

= Uvaria winkleri Diels, Bot. Jahrb. Syst. 38(3): 240 (1907). Type. Cameroon - South-West Region • H. Winkler 1466 (holotype: B (not found, destroyed?)); Moliwe.

= Uvaria megalantha Diels, Bot. Jahrb. Syst. 39(3-4): 472 (1907). Type. Cameroon - South Region • G.A. Zenker 3204 (holotype: B (not found, destroyed?); lectotype: WAG! (WAG0057972), designated by Couvreur et al (2022); isolectotypes: BM! (BM000636652), E! (E00147958), G! (G00412220), GOET! (GOET005733), HBG! (HBG502487), K! (K000198800), L! (L.1768578), M! (M0107939), P! (P01982908), S! (S07-13392), WU! (WU0025787), Z! (Z-000000876, Z-000000877)), Kamerun. Bipinde; 3°05'N, 10°25'E; 1904.

Description.

Tree 3-20 m tall, D.B.H. 2-25 cm; young branches glabrous, old branches glabrous. Petiole 5-21 mm long, 2-6 mm wide, glabrous. Leaf lamina 254-636 mm long, 66-177 mm wide, length:width ratio 2.3-6.2, elliptic to oblong to obovate, coriaceous, base acute to rounded, sometimes truncate or subcordate, apex acute to acuminate, acumen 5-24 mm long; surface above glabrous, surface below glabrous; midrib impressed above, raised below, glabrous above, glabrous below; secondary veins 16-28 pairs, weakly brochidodromous, impressed above, raised below; tertiary veins reticulate. Inflorescences borne on trunk and branches or axillary, composed of 1 flower. Flower pedicel (5) 10-40 mm long, 1.4-4 mm, increasing toward the apex up to 10 mm in diameter, pubescent to glabrous. Flowers bisexual, buds ovoid to globose, pedicellate, 9-22 mm high, 8-30 mm in diameter, pubescent. Bracts 1 to 6, upper bract 4-13 mm long, 6-17 mm wide, ovate, appressed, enclosing the bud, pubescent outside, glabrous inside. Sepals 3, 5-14 mm long, 7-17 mm wide, imbricate to fused at base, puberulent outside, glabrous inside, dull green. Outer petals 3, 14-32 mm long, 12-29 mm wide, length:width ratio 0.95-1.5, broadly ovate to ovate, pubescent to puberulent outside, puberulent to glabrous inside, wine red (cream in immature flowers) outside, wine red (cream in immature flowers) inside. Inner petals 3, 9.5-30 mm long, 8-23 mm wide, length:width ratio 0.9-1.9, broadly ovate to ovate, pubescent to puberulent outside, puberulent to glabrous inside, wine red (cream in immature flowers) outside, wine red (cream in immature flowers) inside. Stamens 1900 to 2500, 2-3.8 mm long, 0.3-0.5 mm wide, anthers linear, connective prolongation truncate. Carpels 5 to 33, 2-6.5 mm long, 0.8-1.5 mm wide, pubescent, free; stigma 0-2 mm long, 1-1.8 mm wide, coiled, pubescent, covered with an exudate at anthesis. Fruiting pedicel 13-31 mm long, 3-8 mm in diameter, glabrous. Monocarps 1 to 10, 22-55 mm long, 17-32 mm wide, length:width ratio 1.3-2, cylindrical to ovoid, longitudinally ridged, sparsely pubescent to glabrous, red to orange when ripe, green when immature; stipe 0-8 mm long, 3-10 mm wide, slightly pubescent to glabrous. Seeds 8-23 per monocarp, biseriate, 13 to 29 mm long, 3-14 mm wide, semicircular.

Distribution.

Endemic to Lower Guinean Domain of the Guineo-Congolian Region: Cameroon, Equatorial Guinea (Bioko Island) and Nigeria.

Habitat and ecology.

Lowland and premontane mature and old secondary rain forests, sometimes in swamp forests. Altitude: 50-1000 m a.sl.

Phenology.

Flowers and fruits collected all year.

Vernacular names.

Cameroon: ‘Ikeinju’ in Bakweri (Mbani 14).

Uses.

The young leaves are eaten (Cheek 5180) and the fruits are used for cough and dye (Mbani 14).

Notes.

This species is easily differentiated from the other species by its petals being wine red on both sides in mature flowers. Apart from this character, Ud. connivens resemble Ud. calophyllum and Ud. fuscum in having great leaves (more than 25 cm long). It can be distinguished from these species by being glabrous when young and old, and by having a distinct flower pedicel 10-40 mm long, whereas Ud. calophyllum and Ud. fuscum flowers are subsessile (flower pedicel 10-15 mm maximum in Ud. fuscum var. magnificum ).

Preliminary conservation status.

This species has been assessed previously as Near Threatened NT but the assessment needs to be updated ( Peguy 1998). Here, the EOO of this species is estimated at 107,423 km2 and its AOO at 268 km2. Based solely on AOO value, it would qualify for Endangered, but none of the other B2 sub-criterion are met. It is relatively widespread in Cameroon with more than 10 locations, thus no longer qualifies for Near Threatened NT. Following IUCN criterion B, and it is assigned a preliminary updated conservation status of Least Concern LC.

Additional specimens examined.

Cameroon - Littoral • T.L.P. Couvreur 620 (MPU, YA), Ebo Wildlife Reserve, Djuma permanent camp. On Djuma-Djuma trail; 4°20'23.59"N, 10°14'41.58"E; alt. 335 m; 14 Feb. 2014 - South Region • G.A. Zenker 2624 (B, BM, G, K, L, M, MA, P, P, WAG), Bipindi ; 3°05'N, 10°25'E; 1903 • G.A. Zenker 3401 (BM, G, K), Bipinde; 3°05'N, 10°25'E; 1907 • G.A. Zenker 3487 (BM, G, K), Bipinde; 3°05'N, 10°25'E; 1908 • G.A. Zenker 358 (B, G, M, P, P, U, WAG), Mimfia; 3°04'N, 10°23'E; Sep. 1913 • G.A. Zenker 3845 (BM, K), Bipindi; 3°05'N, 10°25'E; 1909 • J.J. Bos 5412 (P, WAG), 6 km S. of Kribi, 2-4 km E. of Grand Batanga road; 2°53'N, 9°55'E; 26 Sep. 1969 • T.L.P. Couvreur 383 (MPU, WAG, YA); Océan, Campo Ma an National Park , 5 km after main entrance; 2°21'19.48"N, 10°15'33.59"E; alt. 300 m; 15 Feb. 2012 • T.L.P. Couvreur 484 (MPU, YA), 15 km east from Lele village ; 2°16'39.39"N, 13°17'37.19"E; alt. 549 m; 09 Sep. 2013 • T.L.P. Couvreur 706 (MPU, WAG, YA), Campo Ma'an National Park, 11 km on trail from Ebinanemeyong village, on road, 7 km from Nyabessan to Campo town ; 2°28'25.42"N, 10°20'39.1"E; 14 Feb. 2015 - South-West Region • B.-A. Nkongmeneck 959 (YA); Fako, Idenau, Mt Cameroun, versabt de Idenao. Feuille IGN 1/200 000 Buea / Douala ; 4°12'N, 9°05'E; alt. 300 m; 23 Feb. 1985 • C.F. Tekwe 87 (K, SCA, YA), Isobi, above Isobe ; 4°10'N, 9°00'E; alt. 40 m; 10 Jun. 1992 • D. Kenfack 1507 (MO), Mokoko ; 4°27'N, 9°04'00.12"E; 24 Apr. 2001 • D. Kenfack 879 (MO); Ndian, Korup National Park, 11 km from Mundemba, along Fabe road; 5°05'03.43"N, 9°32'20.63"E; alt. 9 m; 11 Jul. 1997 • D.W. Thomas 2264 (K, MO, P, WAG, YA); Ndian, Map # NB 32 IV Buea-Douala. South Korup Reserve ; 4°55'N, 8°50'E; alt. 50 m; Jul. 1983 • D.W. Thomas 4447 (K, MO, P, YA), forest and meadows on the gently sloping side of Mt Cameroun above small Koto village ; 4°18'N, 9°06'E; alt. 550 m; 06 Mar. 1985 • D.W. Thomas 5533 (P, YA), forest between Kindonge and small Ekombe, Southern Bakundu Forest Reserve ; 4°35'N, 9°23'E; alt. 200 m; 10 Feb. 1986 • D.W. Thomas 5537 (P, YA), at Bonenza , 2 km N of Limbe-Idenao road; 4°03'N, 9°05'E; alt. 300 m; 10 Feb. 1986 • D.W. Thomas 6928 (P, YA), footpath toward Matene from Mbilishe; 6°14'N, 9°26'E; alt. 400 m; 1987 • D.W. Thomas 7961 (MO, P, YA), Steep hillside south of Esukutang village ; 5°23'N, 9°00'E; alt. 300 m; 25 May. 1988 • D.W. Thomas 9875 (K, SCA, YA), West of the Onge River and ridges on ' Thump Mount' ; 4°20'N, 8°57'E; alt. 200 m; 09 Nov. 1993 • F. Nguembock 76 (K); Fako, Mabeta-Moliwe Reserve ; 3°58'N, 9°14'E; 10 Mar. 1992 • G. Mann 763 (BM, K), Ambas Bay ; 4°01'N, 9°12'E; Feb. 1861 • G.K. Gottsberger 130307/21 (ULM, WAG), c. 300 m from Banyang Mbo Research Station ; 5°08'N, 9°30'E; 13 Mar. 2007 • G.P. Tchouto Mbatchou 685 (K, SCA, YA), Bomana secondary forest. Transect OA, Plot OA0Z; 4°15'N, 9°01'E; alt. 200 m; 05 Oct. 1993 • I. von Rege 87 (K, SCA, YA); Fako, Mabeta , 6 km SE Limbe SBL; 3°59'N, 9°17'E; alt. 60 m; 11 Aug. 1993 • J. Bongyu 73 (K); Fako, Mabeta-Moliwe Reserve ; 3°58'N, 9°14'E; 06 Apr. 1992 • J. Nemba 550 (MO, P, YA), Secondary growth and old growth forest along Kumba-Mamfe road at mile 14 Lkiliwindi ; 4°44'N, 9°29'E; alt. 200 m; 15 Jun. 1987 • J. Nemba 56 (L, P, U, YA), 5 kms west of Kumba-Mamfe road near Konye ; 4°55'N, 9°36'E; alt. 300 m; 25 Mar. 1986 • J.-P. Ghogue 1551 (YA), Bimbia Bonadikombo (former Mabeta Moliwe ), 18 km SE Limbe; 3°59'36"N, 9°15'43"E; alt. 50 m; 10 Apr. 2003 • J.F. Villiers 2483 (P, YA), 4 km E Bomana , 34 NW limbé; 4°12'55.22"N, 9°06'11.21"E; 14 Dec. 1984 • J.I. Wheatley 194 (K, SCA, YA); Fako , Mabeta-Moliwe TC 10; 4°01'N, 9°16'E; alt. 50 m; 20 Apr. 1992 • J.I. Wheatley 326 (K, SCA, YA); Fako , Mabeta-Moliwe TD 5835 m; 4°01'N, 9°16'E; alt. 100 m; 24 Jun. 1992 • J.J. Wieringa 45 (WAG); Fako, Limbe, Bakingini , forest above 'mile 11 village'; 4°04'10"N, 9° 03'50"E; alt. 160 m; 20 Jan. 1994 • J.J. Wieringa 5839 (BR, FHO, MO, WAG), Bakingini, at edge of plantation area above Mile 11; 4°04'10.8"N, 9°03'27.6"E; alt. 150 m; 06 Mar. 2007 • J.M. Dalziel 8243 (K), Buea 1, Buea to Mayuko ; 4°09'N, 9°14'E; 13 Feb. 1927 • J.M. Mbani 14 (K, SCA, YA), Njonji ; 4°08'11.04"N, 8°59'22.56"E; alt. 100 m; 13 Feb. 1992 • J.M. Mbani 382 (K, SCA, YA), Ekumbe Mofako , Plot M 19; 4°28'N, 9°04'E; 21 May. 1994 • J.P. Watts 245 (K, YA), Moliwe, Makota River watershed .TD 5832 m; 4°00'N, 9°15'E; alt. 100 m; 29 Apr. 1992 • J.P. Watts 394 (K, SCA, YA), c. 5 km South East of Moliwe , TF + 4000 m; 4°02'N, 9°17'E; alt. 100 m; 09 Jun. 1992 • J.P. Watts 782 (K, SCA, YA), forest to the West of Onge River, about 4 km West of Liwenyi village (c 14 km North of Idenau ); 4°23'N, 8°59'E; alt. 260 m; 28 Oct. 1993 • M. Etuge 156 (MA, MO, P, WAG, YA), Bakolle Bakossi, on Kumba - Mamfe road; 5°01'N, 9°40'E; alt. 350 m; 24 May. 1986 • M. Etuge 2390 (K, YA), Max's trail, Nyasoso ; 4°49'39.72"N, 9°40'51.96"E; alt. 1100 m; 24 Jun. 1996 • M. Etuge 2396 (K, YA), Max's trail, Nyasoso ; 4°49'39.72"N, 9°40'51.96"E; alt. 1100 m; 24 Jun. 1996 • M. Etuge 6506 (YA), Mungo River F.R., Mungo F.R; 4°44'17"N, 9°33'38"E; 22 Feb. 2006 • M.R. Cheek 5180 (K, P, SCA, YA), Liwenyi, Low altitude forest on the West bank of the Onge river above the first set of rapids, that is about 1-2 hours walk inland from Enyenge. Grid ref. and alt. approx. Local names and uses from Clement Offu ( Enyenge ); 4°17'N, 8°58'E; alt. 50 m; 28 Oct. 1993 • M.R. Cheek 5462 (K, SCA, YA), Low altitude forest above oil palm plantation. Reached after c. 40 minutes walk N then E from Njonji. Hunters path to ' Lake Njonji' . Little farming, but many gaps and fallen trees at low altitude, c. 150-300 m, crossing and running alongside substantial seasonal stream. At c. 400 m, Hypselodelphyus-Aframomum thicket dominates; 4°08'N, 9°01'E; alt. 300 m; 18 Nov. 1993 • M.R. Cheek 8164 (K, YA); Ndian, Korup National Park, Ekundu Kundu , Transect 10, c. 1100 m; 5°08'N, 8°55'E; alt. 170 m; 25 Apr. 1996 • N. Ndam 1076 (K, SCA), Bonjare, Plot 09; 4°26'N, 9°01'E; alt. 220 m; 30 Apr. 1994 • N. Ndam 1118 (K, SCA), Bonjare, Plot 10; 4°26'N, 9°01'E; alt. 270 m; 01 May. 1994 • N. Ndam 708 (K, YA), Bomana-Koto Rd c 500 m Bearing 305deg towards Onge river 3 hr walk from the rd; 4°13'N, 9°04'E; alt. 400 m; 18 Oct. 1993 • P. Nkeng 37 (K, SCA, YA), Etome ; 4°03'N, 9°07'E; alt. 300 m; 02 Mar. 1992 • R.G. Letouzey 15175 (P, YA); Ndian, Rivières Mosongosele et de Ndian depuis Mosongosele jusqu'à l'entrée amont de la mangroce, env. 20 km au SW de Mundemba (feuille IGN 1/200 000 Buea-Douala ); 4°49'27.3"N, 8°45'15.72"E; 13 Jun. 1976 - R.G. Letouzey 15177 (MO, P, P, WAG, YA), Au SW de Mosongosele , 20 km WSW Mendumba; 4°54'N, 8°45'E; 14 Jun. 1976 • S. Cable 2329 (K, YA); Ndian, Korup National Park, Ekundu Kundu, path to Esoki about 4 km from Ekundu-Kundu (as on 27th); 5°03'16.8"N, 8°56'58.5"E; alt. 350 m; 29 Apr. 1996 • S. Cable 611 (K, SCA, YA), Dikulu, Coastal lowland rain forest along Mangrove stream; 3°59'N, 9°14'E; alt. 50 m; 17 Dec. 1993 • S.N. Ekema 1078 (K, YA), Mokoko Forest Reserve, Boa / Likinge ; 4°24'N, 9°00'E; alt. 150 m; 31 May. 1994 • T.C.H. Sunderland 1264 (K, SCA, YA), Mabeta-Moliwe : TB 6000 m; 4°03'N, 9°16'E; alt. 40 m; 22 Apr. 1992 • T.C.H. Sunderland 1536 (K, K, SCA, YA), Nyasoso, Mt Kupe: Max's Trail ; 4°48'05.04"N, 9°42'29.16"E; alt. 1600 m; 09 Jul. 1992 • T.D. Maitland 537 (K, P), Buea area, at Balifamba; 4°10'N, 9°18'E; alt. 731 m; 1929 • T.L.P. Couvreur 1016 (MPU, WAG, YA), Bayang Mbo Wildlife Sanctuary, after Mbu river ; 5°21'26.28"N, 9°30'05.88"E; alt. 253 m; 26 Mar. 2016 • T.L.P. Couvreur 1051 (MPU, WAG, YA), Mt Cameroon National Park, on the Bomona trail, behind Bomona village, 10 km NW from Idenau ; 4°17'48.27"N, 9°04'43.77"E; alt. 690 m; 03 Apr. 2016 • W.G. Gosline 235 (K, YA), Kupe Village, trail to Kupe rock saddle; 4°47'10"N, 9°41'30"E; alt. 950 m; 28 Nov. 1999 • W.J. Baker 294 (K, SCA, YA); Fako, Mabeta , 6 km SE Limbe SBL; 3°59'N, 9°17'E; 10 Aug. 1993 - Unknown major area • W.G. Gosline 209 (K, WAG, YA), Meme Division, Mahole-Bintulu road; 4°47'30"N, 9°36'12"E; alt. 300 m; 24 Nov. 1999 . Equatorial Guinea - Bioko Norte • M.G. Carvalho 4220 (K, MA), BIOCO: Malabo - Cupapa, km 22-23, margenes del rio Ejoa , 32NMK 9005, 200 m; 3°43'12"N, 8°51'36"E; 06 Jul. 1989 - Bioko Sur • W.R.Q. Luke 11955 (EA, K, MA), Moaba - Moka trail, Biadyi River Camp Pt 140; 3°17'06.4"N, 8°38'17.81"E; alt. 650 m; 16 Mar. 2007 • W.R.Q. Luke 13184 (K, MA), Moraka pt 340 to 342; 3°16'13.8"N, 8°28'29.64"E; alt. 3 m; 28 Jan. 2009 . Nigeria - Cross River State • C.F.A. Onochie FHI36080 X (K); Calabar, Oban Group Forest Reserve , between Akor and Orem; 5°36'N, 8°35'E; 21 Jan. 1957 • E.U. Ujor FHI31636 (K); Calabar, Ikot Ewa ; 4°57'24.84"N, 8°38'44.88"E; 05 Jul. 1952 • H.D. Onyeachusim FHI54055 (K); Calabar, between miles 57-58 Osomba village on Calabar-Mamfe road; 5°27'23.32"N, 8°39'55.34"E; 21 Feb. 1964 • M.C. Ejiofor FHI21898 (K); Calabar, Oban Group Forest Reserve ; 5°36'N, 8°35'E; 07 May. 1952 • M.G. Latilo 23 (K); Calabar, Akamkpa Rubber Estate. Calabar River Division ; 5°18'N, 8°20'E; 21 Mar. 1959 • M.G. Latilo 32 (K, MO); Calabar, Akamkpa Rubber Estate. Dukwe felling area. Calabar River Division ; 5°18'N, 8°20'E; 23 Mar. 1959 • P.A. Talbot 1254 (BM, K), Oban; 5°13'23.28"N, 8°33'06.9"E; 1911 • P.A. Talbot 158 (BM); Calabar, Oban; 5°19'N, 8°34'E; 1911 • P.A. Talbot 402 (BM), Oban; 5°13'23.28"N, 8°33'06.9"E; 1911 • P.A. Talbot 404 (BM, K), Oban; 5°13'23.28"N, 8°33'06.9"E; 1911 • P.A. Talbot 433 (BM), Oban; 5°13'23.28"N, 8°33'06.9"E; 1911 • P.A. Talbot 434 (BM), Oban; 5°13'23.28"N, 8°33'06.9"E; 1911 • P.A. Talbot 82 (BM, K), Oban; 5°13'23.28"N, 8°33'06.9"E; 1911 • P.A. Talbot s.n (BM); Calabar, Oban; 5°13'23.28"N, 8°33'06.9"E; 1912 • P.P.C. van Meer 1412 (WAG); Calabar, Oban Group Forest Reserve, East Block ; 5°31'N, 8°41'E; alt. 200 m; 21 Apr. 1971 • P.P.C. van Meer 1430 (U); Calabar, Oban Group Forest Reserve, East Block ; 5°31'N, 8°41'E; alt. 200 m; 22 Apr. 1971 • P.P.C. van Meer 1430 (WAG); Calabar, Oban Group Forest Reserve, East Block ; 5°31'N, 8°41'E; alt. 200 m; 22 Apr. 1971 • R.W.J. Keay FHI28191 (K); Ikom District, Afi River Forest Reserve , near Aboabam , forest by river Nkem ; 6°11'52.16"N, 8°58'40.48"E; 09 Dec. 1950 .

Figures

Figure 7. Uvariodendron connivens (Benth.) R. E. Fr. A leaf B flower, two outer and one inner petal removed C outer petal inner view D stamen, front view. Uvariodendron angustifolium (Engl. & Diels) R. E. Fr. E leaves F flower, semi top view G flower, bottom view showing bracts H flower, bottom view, bracts removed I outer petal, outer view J stamen, front view K carpel, front view and detail of ovules. A-D from Mann 1159 A-K from Vigne 1610. Drawings by Helene Lamourdedieu, Publications Scientifiques du Museum national d'Histoire naturelle, Paris.

Figure 15. Uvariodendron connivens (Benth.) R. E. Fr. A young branch with leaves, upper side B detail of petiole and base of leaf, upper side C fruiting material with unripe monocarp, side view D flower, top view E flower, side view F flower bud, top view G flower bud, bottom view H flower, one sepal removed, semi-bottom view. A, B Couvreur 383 C, F, G Couvreur 620 D, E Couvreur 1016 H Couvreur 1051. Photos Thomas Couvreur.

Figure 16. Distribution map of Uvariodendron connivens. Shades of grey represent elevation, from white (sea level) to darker grey (higher elevation). The inset shows the extent of the map over Africa.