Solmsia calophylla Baill.

Rogers, Zachary S. & Fuentes-Soriano, Sara, 2021, Solmsia Baill.: a taxonomic revision of an endemic New Caledonian genus of Thymelaeaceae, Adansonia (3) 43 (12), pp. 125-150 : 139-148

Solmsia calophylla Baill.

( Figs 1 ; 6 ; 7 )

Adansonia, Recueil d’Observations botaniques 10: 38 ( Baillon 1871) [12.VI.1871].

— Lectotype (designated here by Rogers): New Caledonia. Grande Terre, Province du Sud, Mont-Dore Commune , “Collines ferrugineuses de la Baie du Prony”, [general area: c. 22°19’S, 166°50’E], IX.1868, imm. fr., fr., B. Balansa 263 (lecto-, P [ P00239783 ]!, Fig. 8 ; isolecto-, A [ A01005249 ]!, NY [ NY03101927 ]!, P [ P06622659 ]!) [residual syntypes (probable and possible) indicated in Specimens examined section].

Solmsia chrysophylla Baill. , Adansonia, Recueil d’Observations botaniques 10: 38, 39 ( Baillon 1871) [12.VI.1871]. — Solmsia calophylla Baill. var. chrysophylla (Baill.) Guillaumin, Notulae Systematicae 1: 108, 109 ( Guillaumin 1909) [20.XII.1909]. — Lectotype (designated here by Rogers): New Caledonia. Grande Terre, Province du Sud, Mont-Dore Commune , “Collines ferrugineuses de la Baie du Prony”, [general area: c. 22°19’S, 166°50’E], IX.1868, fl. bud, fr., B. Balansa 262 (lecto-, P [ P06622241 ]!, Fig. 9 ; isolecto-, P[P00239812]!, P[P06622240]!), syn. nov. [residual syntypes (probable and possible) indicated in Specimens examined section].

DISTRIBUTION, PHENOLOGY AND ECOLOGY. — Solmsia calophylla is endemic to the Grande Terre of New Caledonia ( Fig. 2 ), where it is restricted to the “maquis” zone, a diverse vegetation type associated with ultramafic substrates most common in the southern quarter of the island (Massif du Sud), but also with numerous relatively small subcoastal outcrops distributed sporadically northwards along the NW coast. The maquis, comprising less than one-third of New Caledonia (c. 5500 km 2), is composed of low, shrubby vegetation that grows on several different variants of ferralsols (also known as oxisols), which are relatively nutrient poor and often rich in heavy metals, especially nickel (region locally known as “maquis minier”) ( ORSTOM 1981; L’Huillier et al. 2010; Isnard et al. 2016). Most populations occur in the southern half of the expansive Massif du Sud region. Disjunct ultramafic outcrops host populations at several sites (running from S to N): Massif Mé Maoya, Massif du Boulinda, Plateau de Tiéa, Massif de Koniambo, Mont Oua Tilou, Massif de Taom/Mont Ouazangou.

Solmsia calophylla occurs from near sea level up to 1000 m elevation, but most herbarium collections were made from within the elevation range of 100 to 600 m. The highest known populations are found around Montagne des Sources, Massif du Humboldt, Col de Vulcain, Mont Koniambo and Mont Taom. The species is locally common at some sites, particularly in the south, and plants have adapted to a wide variety of habitats. Populations occur in several different subtypes of maquis, ranging from the more widespread, open, eroded upland maquis, where plants are exposed to higher light levels, winds, and poorer-quality soils (Morphogroup B, “chrysophylla form”), to the hydromorphic maquis associated with low-lying watercourses on colluvio-alluvial substrates that yield a more protected, low forest maquis vegetation, with much richer soils (Morphogroup A, “calophylla form”). For more specific ecological details and edaphic associations see the corresponding subsection above within the Results and discussion text.

Flowering and fruiting may take place year round, and sometimes the same plant will have floral buds and mature, dehisced fruits simultaneously present on a single branch. New Caledonia’s climate has been classified into four seasons (Mueller-Dombois & Fosberg 1998), and rain received during the two intervening wetter seasons significantly impact flowering activity in Solmsia . The most frequent month for flowering recorded by far is March, near the end of the warm, rainy season (perhaps due to collection bias), followed by a second less prolific peak in July during the middle of a season of moderate precipitation. The most commonly reported fruiting period for S. calophylla is June through November.

Flowers were reported as fragrant on the labels of five collections (all staminate). Pistillate plants in flower are frequented by ants (MacKee 43461, MO!, NOU!, P!; also see a separate visitation event in photo, Fig. 7E ). It would appear that pollen is the reward because flowers of both sexes lack potentially nectar-producing organs such as petaloid organs and a subgynoecial disk. Ant visitors have also been observed carrying seeds away from the parent plant in the closely related Lethedon on Grande Terre (J. Munzinger, pers. comm.). Besides the possibilities of intentional or unintentional pollination, it seems likely that ants in both genera are drawn to the thin, outer epidermis of the seeds that include a semi-fleshy arillate appendage at one end ( Fig. 7F ).

VERNACULAR NAME. — The name “Mouoguéporo” (Bourail) has been applied to Solmsia calophylla var. chrysophylla (fide Guillaumin 1911). No specimen vouchers documenting that local name or any other common name for Solmsia have been located in herbaria or literature.

USES. — No uses have been reported for Solmsia calophylla , but the species would make an attractive ornamental if brought into cultivation ( Figs 6 ; 7 ), especially in regards to those individuals with persistent golden velutinous pubescence on their abaxial leaf surfaces (i.e., Morphogroup B, “chrysophylla form”).

SPECIMENS EXAMINED [crosses (+): Morphogroup A, “calophylla form”; squares (□): Morphogroup B, “chrysophylla form”; asterisks (*): morphological intermediates]. — New Caledonia. Grande Terre, Province du Nord, Canala , [sheets variously annotated: “ Kanala, Mt. Dore ” or “Kanala, etc.”], [Canala: c. 21°31’S, 165°57’E], 1861-1867, fl. bud, fl., imm. fr., E. Vieillard “145, 146” (F!, G!, GH!, K!, P[4 sheets]!, Z!) [possible residual syntypes of Solmsia chrysophylla ] [□] ; [sheet annotated: “ Montagnes ferrug. Kanala 8” [final letter or number illegible], [ Canala : c. 21°31’S, 165°57’E], 1861-1867, fl., E. Vieillard “145=146” (P!) [possible residual syntype of Solmsia calophylla (upper-right branch), and possible residual syntypes of Solmsia chrysophylla (two left-hand branches)] [+ and □] ; [sheets annotated: “ Bois des Montagnes Kanala ”], [ Canala : c. 21°31’S, 165°57’E], 1855-1860, fl. bud, E. Vieillard 146 (P[3 sheets]!, Z!) [possible residual syntypes of Solmsia chrysophylla ] [□] ; [sheet annotated: “ Bois des Montagnes Kanala ”, but locality questionably within distribution of Morphogroup A, possibly instead collected near Mont-Dore, see Results and discussion section], [ Canala : c. 21°31’S, 165°57’E], 1855-1860 [a few sheets with typewritten labels dated 1861-1867], fl., imm. fr., E. Vieillard 147 (GH!, K!, P[5 sheets]!) [probable residual syntypes of Solmsia calophylla ] [+] ; Canala , dans les terrains ferrugineux, [21°31’S, 165°57’E], XI.1869, fl., imm. fr., fr., B. Balansa 1905 (G!, P[4 sheets]!) [possible residual syntypes of Solmsia chrysophylla ] [□] ; Canala, [21°31’S, 165°57’E], 20.II.1912, fl. bud, fl., F. Sarasin 553 (Z!) [□]. — Kaala-Gomen, Mont Ouazangou , pente ouest, maquis sur terrains, 300-500 m, [20°45’S, 164°29’E], 4.III.1972, fl., H. S. MacKee 25117 (BM!, CANB!, K!, L!, MO!, NOU!, P!) [□ and *] ; Koniambo, maquis ligno-arbustif, Relevé TK1, [21°01’S, 164°44’23”E], 19.IX.2003, fl. bud, fl., J. Fambart-Tinel & F. Roumagnac 40 (NOU!, P!) [□] ; Mont Ouazangou , pente ouest, maquis, pente rocheuse serpentineuse, 700 m, [20°45’S, 164°29’E], 26. III.1980, fl., H. S. MacKee 37962 (L!, NOU!, P!) [□] ; Mont Taom (contrefort nord-ouest), maquis, terrain serpentineux, 900 m, [20°47’S, 164°35’E], 8.II.1980, fl., H. S. MacKee 37842 (CANB!, G!, K!, L!, MO!, NOU!, P!) [□]. — Koné, Koniambo, Koné , [21°01’S, 164°48’E], 18.I.1925, fr., A. U. Däniker 958 (Z!) [□] ; Massif de Koniambo , south of Voh, 750 m, [21°00’S, 164°49’E], 6.I.1983, fl. bud, G. McPherson 5300 (MO!) [□] ; Massif de Koniambo , rocky hillside with maquis, 650 m, [21°00’S, 164°45’E], 25.III.1987, fl., K. L. Wilson 7153 (NSW!) [□] ; Pouembout, Tiéa (plateau), maquis dégradé, terrain serpentineux altéré, 350-450 m, [21°11’S, 164°54’E], 27.XII.1974, fl. bud, fl., H. S. MacKee 29572 (G!, K!, L!, MA, MO!, P!) [□]. — Kouaoua, Kouaoua mine SLN, [21°23’00”S, 165°48’40”E], 14.II.1978, fl. bud, fl., P. Cabalion 405 (MO!, P!) [□] ; Kouaoua, Ouaseoua , maquis dégradé, alluvions serpentineuses, 30 m, [21°24’S, 165°46’E], 29.III.1977, fl. bud, fl., H. S. MacKee 32967 (NOU!, P!) [□]. — Pouébo?, [sheet annotated: “ Montagnes de Pouébo ” and bearing specimens belonging to both morphogroups, the locality is doubtful for both groups because the site is an extremely disjunct northern population that overlies non-ultramafic substrate, see Results and discussion section], [Pouébo: c. 20°24’S, 164°34’E], 1855-1860, fl., E. Vieillard 254 (P!) [possible residual syntypes of Solmsia calophylla (three leftmost branches), and possible residual syntype of Solmsia chrysophylla (rightmost branch)] [+ and □]. — Poya (northern), Massif du Boulinda , 500 m, [21°16’S, 165°08’E], 28.VIII.1987, fl., imm. fr., T. Jaffré 2881 (MO!, NOU!, NSW!, P!) [□] ; Massif du Boulinda, maquis, 400 m, [21°16’S, 165°08’E], 23.II.1978, fl. bud, fl., fr., P. Morat 5944 (NOU!, P!) [□] ; Massif du Boulinda , route de la mine St-Louis, au niveau du petit creek, en maquis minier, [21°16’S, 165°08’E], 22.II.1978, fl. bud, fl., fr., B. Suprin 251 (NOU!) [□]. — Voh, Oua Tilou , sur serpentine, 800 m, [20°51’57”S, 164°51’28”E], 14.IV.1951, st., A. Guillaumin 12406 (G!, P!, Z!) [□]. — Province du Nord/Sud, Bourail , crest to N of Col des Roussettes , between upper valley of Houaïlou and Azareu (spur of Mé Maoya Massif), 700 m, [21°26’S, 165°27’E], 22.I.1963, fl. bud, fl., H. S. MacKee 10103 (K!, L!, P!)[□] ; entre Thio et Houaïlou , 1910, st., M. Fetscherin s.n. (P!) [□]. — Province du Sud, Boulouparis , Col de Vulcain , serpentine scrub, 900 m, [21°54’S, 166°23’E], 11.XI.1950, imm. fr., M. G. Baumann-Bodenheim 8123 (K!, P!, Z!) [□] ; Bwa Bwi, contrefort Sud-Est, maquis arbustif, substrat ultramafique, 890 m, 21°46’40.8”S, 166°17’57.3”E, 23.XI.2016, imm. fr., D. Bruy & J. Munzinger 528 (MPU, NOU, P!) [+] ; contrefort Nord du Koungouhaou Nord, sous la Concession Byzance Red , maquis haut, [21°47’30”S, 166°08’00”E], 12.I.2005, fl. bud, J. Munzinger & G. Dagostini 2634 (MO!, NOU!) [□] ; La Ouaménie , terre ferrugineuse, 600 m, VIII.1881, fl., imm. fr., A. Brousmiche s.n. (P!) [□ and *] ; Massif du Humboldt , growing in low forest, 800 m, [21°53’S, 166°25’E], 14.XI.1982, imm. fr., fr., W. G. Ziarnik 75 (BRI!, MO!, NOU!) [□] ; bassin de la Tontouta, route du Humboldt, maquis dominé par Gynostoma chamaecyparis, sol érodé sur péridotites serpentinisées, [22°00’S, 166°09’E], 21.III.1996, fl., T. Jaffré et al. 3301 (NOU!, P!) [□] ; Mine Galliéni , at upper end of Tontouta River valley, partially disturbed maquis, near remnant forest, 740 m, 21°54’32”S, 166°21’30”E, 24.IX.1998, imm. fr., P. Lowry 5085 (MO!, NOU!, P!) [□] ; Mont Do, upper region towards top, depauperate bush and heath on E slope, rather dense and humid Araucaria montana forest on W slope, serpentine and peridotite, 700-1014 m, [21°45’S, 166°00’E], 28.XI.1966, imm. fr., F. Ehrendorfer 6600-138.11 (W, WU!) [□] ; Mount Gouavi , south slope, above the lower Tontouta Valley , 400 m, 21°56’S, 166°12’E, 10.II.1991, fl. buds, A. M. Buchanan 11945 (HO, MEL!) [□]. — Boulouparis/Païta, bord de la Rivière du Humboldt , [21°57’S, 166°22’E], 15.XII.1964, fl. bud, J. P. Blanchon 1310 (NOU!) [□]. — Dumbéa, Dumbéa , [22°09’S, 166°27’E], 1910, fl. bud, A. D’Alleizette 507 (P!) [□] ; bord de la Dumbéa , [22°09’S, 166°27’E], 1911, fl. bud, I. Franc 27 (G!, P!) [□] ; North Dumbéa Valley , between 1 st and 4 th bunkers of abandoned mine, 400-800 m, 13. XI.1955, fr., H. S. MacKee 3350 (K!, P!, US!) [□] ; Barrage de Dumbéa , 110 m, [22°08’15”S, 166°31’30”E], 11.XI.2007, fr., J. Munzinger et al. 4666 (NOU!, P!) [□] ; Mine Werguin (haute Dumbéa), 200 m, [22°09’S, 166°27’E], 30.IX.1939, fl. bud, fl., R. Virot 212bis (A!, P!) [□] ; Montagne des Sources , [22°07’S, 166°36’E], 24.I.1950, fl. bud, L. Baas-Becking 5966 (G!, Z!) [□] ; Montagne des Sources , 900-1000 m, [22°07’S, 166°36’E], 1. IV.1968, fl. bud, L. Bernardi 12483 (G!, K!, L!, P!, Z!) [+] ; Montagne des Sources , mountain plateau, evergreen forest with Araucaria on peridotite, bush and pioneer vegetation on wind exposed slopes and ridges, 600-750 m, [22°07’S, 166°36’E], 30.XI.1966, imm. fr., F. Ehrendorfer 6600-146.32 (W, WU!) [+] ; Montagne des Sources , middle pt, [22°07’S, 166°36’E], 14.VIII.1981, fl., imm. fr., B. C. Stone 14949 (NOU!) [+] ; road to Montagne des Sources , c. 20 km from Nouméa, maquis scrub vegation, secondary growth, 200 m, [22°13’S, 166°36’E], 23.VI.1977, fl., imm. fr., M. Fallen et al. 117 (BISH!, L!, MO!, NOU!, P!) [□] ; road from Nouméa to Montagne des Sources , 14 miles [= 22.5 km] from Nouméa, serpentine area, 60-90 m, 1.VIII.1952, imm. fr., C. McMillan 5196 (A!, L!, P!) [□] ; c. 15 km N of Nouméa, on road to Montagne des Sources , in the major watershed area for the island, 350 m, 14.VIII.1981, imm. fr., D. Mueller-Dombois 81081403 (BISH!) [+] ; versant W de la crête S du Pic Buse , forêt mésophile sur serpentine, 570 m, [22°09’S, 166°35’30”E], 30.XII.1950, st., H. Hürlimann 524 (A!, NY!, P!, Z!) [+] ; pentes sud du Pic Buse ( Haute Boulari ), 600 m, [22°09’S, 166°35’30”E], 18.VI.1939, fl. bud, R. Virot 101bis (A!, P[2 sheets]!) [+]. — Dumbéa?, road to Montagne des Sources , 800 m, [22°07’S, 166°36’E], 13.III.1955, fl. bud, fl., H. S. MacKee 2206 (US!) [+]. — Mont-Dore , Baie de Prony , collines ferrugineuses, [22°19’S, 166°50’E], IX.1868, fl. bud, fr., B. Balansa 262 (P[3 sheets]! [lectotype collection of S. chrysophylla ] [□] ; Baie de Prony , collines ferrugineuses, [22°19’S, 166°50’E], IX.1868, imm. fr., fr., B. Balansa 263 (A!, NY!, P[2 sheets]!) [lectotype collection of S. calophylla ] [+] ; à l’ouest de la Baie des Pirogues , pente raide et rocailleuse, maquis serpentineux bas et épars, 100 m, [22°19’S, 166°41’E], 16.XII.1950, fl. bud, imm. fr., H. Hürlimann 361 (A!, NY!, P!, RSA!, Z!) [□] ; Rivière des Pirogues, W side, near Baie des Pirogues , mangrove swamp and nearby cliffs and beaches, [22°19’S, 166°41’E], 23.V.1977, fl., L. J. Musselman et al. 5355 (NOU!) [□] ; between the N’Go Bay [= Baie Ngo ] and Touaourou , [22°18’10”S, 166°43’E], 1903, fl. bud, O. Rorhdorf 141 (Z!) [□] ; between the N’Go Bay [= Baie Ngo ] and Touaourou , [22°18’10”S, 166°43’E], VIII.1903 - XII.1903, fr., O. Rorhdorf 191 (Z!) [+] ; Baie du Carénage , surrounding hillsides, serpentine area, 60-90 m, [22°18’10”S, 166°50’50”E], 22.VII.1952, fl., imm. fr., C. McMillan 5135 (A!, K!, L!, P!) [+] ; Col de Mouirange, avant le Col, sur la route à Yaté, la forêt mésophile sur serpentine, 200 m, [22°13’30”S, 166°39’15”E], 26.I.1951, fr., H. Hürlimann 751 (A!, NY!, P!, Z!) [□] ; Col de Mouirange , route de Yaté, maquis sur terrain serpentineux, 200 m, [22°12’30”S, 166°40’45”E], 31.I.1969, fl. bud, fl., H. S. MacKee 20233 (BM!, CANB!, G!, K!, L!, MA, MO!, NOU!, NY!, P!, Z!) [□] ; along Nouméa road, along creek, c. 1 km E of Col de Mouirange , low forest, 150 m, [22°12’15”S, 166°42’00”E], 16.III.1981, fl. bud, G. McPherson 3597 (MO!, P!) [+] ; Col de Mouirange , formation paraforestière, ultramafique, [22°12’30”S, 166°40’45”E], 4.III.2006, fl. bud, fl., Y. Pillon et al. 318 (MO!, NOU!) [□] ; Col de Plum ( Mont-Dore ), sur serpentine, 200 m, [22°15’S, 166°36’E], 22.VIII.1950, st., M. G. Baumann-Bodenheim 5613 (Z!) [□] ; Col de Plum, on RT2, along the trail leading to the peak Gué Xi, vegetation is maquis minier, dominated by Soulamea pancheri , Solmsia calophylla , Codia spp. , and Hibbertia spp. , soil is red-brown ultrabasic laterite, 100-450 m, 22°14’48”S, 166°37’26”E, 12.XII.2000, fl. bud, fl., J. W. Horn 3519 (DUKE!) [□] ; La Coulée , [22°14’S, 166°34’E], 12.III.1950, fl., L. Baas-Becking 6004 (Z!) [□] ; Haute Boulari, ravin boisé, terrain serpentineux, 500 m, [22°09’10”S, 166°35’30”E], 28.VI.1978, imm. fr., H. S. MacKee 35307 (K!, L!, MO!, NOU!, P!) [+] ; La Coulée River (Boulari) Valley, 75 m, [22°14’S, 166°34’E], 10. IX.1983, imm. fr., G. McPherson 5776 (MO!) [□ and *] ; La Coulée Valley , NE of Nouméa, secondary scrub near river, [22°14’S, 166°34’E], 30.IX.1983, fr., G. McPherson 5819 (BRI!, MO!, NOU!, P!, PTBG!) [□ and *] ; au-dessus de l’Hôtel des Brugerès, à la Coulée Boulari, collines serpentineuses, 200 m, [22°14’S, 166°34’E], V.1938, fr., R. Virot s.n. (A!, P!) [□] ; Mont-Dore , scrubby woods on serpentine along Rivière La Coulée , 6.5 km upstream from La Coulée, [22°16’S, 166°35’E], 4.VIII.1968, fl. bud, fl., G. L. Webster & R. Hildreth 14535 (BISH!, GH!, NOU!, NSW!, P!) [□] ; Prony, île Casy , 0-40 m, [22°21’15”S, 166°50’35”E], 15.IV.1974, fl. bud, fl., H. S. MacKee 28501 (P!) [□] ; Les Dalmates, route Nouméa-Yaté , terrain serpentineux, maquis dégradé, 150 m, [22°13’S, 166°41’E], 8.III.1969, fl., H. S. MacKee 20271 (BM!, CANB!, G[2 sheets]!, K!, L!, MA, MO!, NOU!, NY!, P!, Z!) [□ and *] ; sur le versant Est du Mont-Dore , forêt à Araucaria , 500 m, [22°16’S, 166°35’E], 21.III.1951, st., A. Guillaumin & M. G. Baumann-Bodenheim 11345 (P!, Z!) [□] ; secteur du Mont-Dore, [22°16’S, 166°35’E], 4.I.1978, fl. bud, fl., imm. fr., T. Jaffré 2130 (MO!, NOU!, P!) [□] ; Mt D’or [= Mont-Dore ], [22°19’S, 166°47’E], 1860, imm. fr., J. A. Pancher s.n. (P[2 sheets]!) [possible residual syntypes of Solmsia chrysophylla ] [□] ; Mont-Dore [one duplicate, P06622211, with the Mont-Dore deleted and replaced with “Kanala” (= Canala)], [Mont-Dore: c. 22°19’S, 166°47’E], [most sheets dated 1855-1860, but with a few alternatively dated 1861, or 1861-1867], fl. bud, fl., fr., E. Vieillard 145 (P[5 sheets]!) [possible residual syntypes of Solmsia chrysophylla , except the small lower-middle branch on P06622209, which is a possible residual syntype of Solmsia calophylla ] [+ and □] ; c. 2 km NW of RT2 and Mont-Dore Road , 19.VII.1977, fl., imm. fr., T.Whaite & J.Whaite 3643 (NSW!) [□] ; Val des Pins supérieur, Pic du Pin , [22°15’00”S, 166°48’58”E], 1.IV.1951, fl. bud, fl., A. Guillaumin & M. G. Baumann-Bodenheim 11924 (L!, P!, Z!) [+] ; Plaine du Lac en 8, route du Carénage , serpentines, sol à tendance hydromorphe, [22°18’S, 166°50’E], 1964, fr., J. P. Blanchon 1168 (L!, NOU!, P!) [+] ; Plaine des Lacs, La Capture , S of Chutes de la Madeleine , c. 6 km from turnoff to entrance of reserve, maquis along creek, just W of road, 280 m, 22°16’01”S, 166°49’19”E, 24.I.2010, fl. bud, fr., P. Lowry et al. 7210 (MO!, P!) [+] ; Plaine des Lacs, partially degraded remnant forest and adjacent maquis W of Camp Penamax , ultramafic substrate, 210 m, 22°16’15”S, 166°49’10”E, 24.I.2010, fl. bud, fl., P. Lowry et al. 7222 (MO!, P!) [□] ; Port-Boisé , [22°21’S, 166°58’E], 6.VI.1977, imm. fr., J. Pusset 17 (NOU!) [+] ; Port- Boisé , [22°21’S, 166°58’E], 16.VII.1977, B. Suprin 170 (NOU!) [+] ; vallée de la Poueta Koure , sur serpentine, [22°19’S, 166°43’E], 19.II.1951, st., M. G. Baumann-Bodenheim 10665 (NY!, P!, Z!) [□] ; vallée supérieure de la Poueta Koure, forêt sur serpentine, 200 m, [22°19’S, 166°43’E], 10.V.1951, fl. bud, A. Guillaumin & M. G. Baumann-Bodenheim 13216 (A!, L!, Z!) [□] ; 5 km au NW de Prony, sur serpentine, 400 m, [22°19’S, 166°49’E], 17.IX.1950, imm. fr., L. Baas-Becking 6081 (L!, P!, Z!) [+] ; Prony, terrain ferrugineux, Plateau Ouest , 100 m, [22°19’S, 166°49’E], VIII.1903, fl., fr., L. Cribs 1545 (P!) [+] ; Prony, terrain ferrugineux, [22°19’S, 166°49’E], XI.1903, fl., fr., L. Cribs 1615 (P[2 sheets]!) [□] ; Prony, localité Bergerie , terrain ferrugineux, 3 m, [22°19’S, 166°49’E], XII.1903, fl. bud, imm. fr., L. Cribs 1728 (P[2 sheets]!) [□] ; Prony, serpentine, [22°19’S, 166°49’E], 5.II.1926, fl. bud, fl., A. U. Däniker 2792 (P!, Z!) [+] ; Prony, littoral, lieux arides, [22°19’S, 166°49’E], I.1907 - II.1914, fl. bud, fl., I. Franc 230A (A!, BRI!, G!, MEL!, NY!, P[2 sheets]!, Z!) [□] ; Prony, littoral, [22°19’S, 166°49’E], I.1914, fl. bud, fl., I. Franc 232 (A!, BRI!, NY!, P!, Z!) [+] ; Prony, terrains arides, [22°19’S, 166°49’E], I.1907, fl. bud, fl., I. Franc 232A (P[2 sheets]!) [+] ; Prony, du littoral, I.1914, fl. bud, fl., I. Franc 234 (G!, NY!) [+] ; Prony, lande et vallées, [22°19’S, 166°49’E], II.1914, fl. bud, I. Franc 1724A (A!, G!, K!, P!) [□] ; route de Prony, [22°19’S, 166°49’E], 24.XII.1977, fl. bud, T. Jaffré 2104 (NOU!) [□] ; Prony, zone maritime, [22°19’S, 166°49’E], 1900-1910, imm. fr., A.- J. Le Rat 1721 (P!) [+] ; Prony, [22°19’S, 166°49’E], 1900- 1910, fl. bud, imm. fr., A.- J. Le Rat 220 (A!, P!) [□] ; c. 10 km E from the junction of Rivière Bleue along Rt. 2, roadside forest on ridge, [22°18’S, 166°50’E], 1.III.1992, fl. bud, fl., H. Koyama & H. Setoguchi 8212 (A!) [+] ; dans la vallée supérieure de la Rivière des Pirogues, maquis serpentineux, [22°19’S, 166°41’E], 29.III.1951, fl. bud, A. Guillaumin & M. G. Baumann-Bodenheim 11527 (Z!) [□] ; ibid., st., A. Guillaumin & M. G. Baumann-Bodenheim 11594 (A!, NY!, P!, Z!) [+] ; ibid., fl. bud, fl., A. Guillaumin & M. G. Baumann-Bodenheim 11611 (L!, P!, Z!) [□] ; ibid., st., A. Guillaumin & M. G. Baumann-Bodenheim 11613 (P!, Z!) [+] ; Rivière des Pirogues, [22°19’S, 166°41’E], 24.X.1923, fr., C. T.White 2241 (A!, BM!, BRI!, P!) [+] ; au NE de St. Louis, collines argilo-ferrugineuses, [22°12’54”S, 166°34’19”E], 15.I.1869, fl. bud, fr., B. Balansa 1191 (A!, BM!, G!, K!, NY!, P[4 sheets]!, Z!) [possible residual syntypes of Solmsia chrysophylla ] [□]. — Païta, Mont Dzumac , [22°03’S, 166°28’E], 8.VII.1965, st., A. Aubréville & Heine 252 (P!) [+] ; Mont Dzumac, pentes, [22°03’S, 166°28’E], VII.1906, fl. bud, I. Franc 230 (P!) [□] ; Mont Dzumac , [22°03’S, 166°28’E], I.1906, fl. bud, I. Franc 230[B] (FI!) [□] ; Mont Dzumac , pentes, [22°03’S, 166°28’E], II.1906, fl., I. Franc 230[C] (FI!) [□] ; Mont Dzumac , pentes, Couvelée , 500 m, [22°03’S, 166°28’E], 15. VIII.1930, fl. bud, fl., imm. fr., I. Franc 2481 (A!, BM!, BRI!, L!, NSW!, NY!, P!, US!, Z!) [□] ; Mont Dzumac , lower slopes, 600 m, [22°03’S, 166°28’E], 9.VIII.1971, fl. bud, B. C. Stone 14805 (BISH!) [□] ; Dzumac , sentier, 600 m, [22°03’S, 166°28’E], 4. IX.1940, fl. bud, imm. fr., R. Virot 171 (A!, P!) [□] ; Mont Mou , [22°04’S, 166°21’E], VII.1906, fl., I. Franc 231 (NY!, P!) [□] ; Mont Mou , 30 km NW of Nouméa, in macchia in rainforest on ultrabasic soil, common in scrub vegetation, 500 m, [22°04’S, 166°21’E], 8.VIII.1981, st., D. Mueller-Dombois 81080805 (BISH!) [□ and *] ; Mont Ni, sur les pentes, collines argilo-ferrugineuses, [21°56’S, 166°25’E], 9.III.1869, fl., B. Balansa 1191a (P!) [possible residual syntype of Solmsia chrysophylla ] [□] ; above a tributary of the Rivière Ni , forested slopes, 820-950 m, 21°59’30”S, 166°29’32”E, 11.XI.2003, imm. fr., fr., G. McPherson & A. Mouly 19221 (MO!, NOU!, P!) [□ and *]. — Païta / Dumbéa , Mts. Kouvele [= Couvelée ] moyens, forêt mésophile sur serpentine, [22°04’S, 166°26’E], 9.V.1951, st., A. Guillaumin & M. G. Baumann-Bodenheim 13106 (P!, RSA!, Z!) [□]. — Thio, Col de Petchicara , [21°35’S, 166°05’E], 3.XII.1986, fl. bud, T. Jaffré 2762 (MO!, NOU!, P!) [□] ; Riv. Koum , [21°43’S, 166°22’E], [no date], J. Munzinger 1012 (MO!, P!) [□] ; Mont Ninga , fourré maquis, 1000 m, [21°45’S, 166°08’E], 19.VI.1975, imm. fr., M. Schmid 5349 (NOU!) [□] ; auf den Hügeln am Ngoye [= on the hills at Ngoye ], 50 m, [21°49’10”S, 166°28’10”E], 29.XI.1902, imm. fr., F. R. R. Rudolf Schlechter 15138 (BM!, G!, K!, L!, P!, Z!) [□] ; à l’embouchure du Tio [= Thio], base des montagnes ferrugineuses, [21°36’46”S, 166°13’00”E], V.1872, fl., B. Balansa 3453 (A!, BM!, G!, K!, NY!, P[3 sheets]!, Z!) [□]. — Yaté, Plaine du Bidou Rouge (Plaine des Lacs), 150 m, [22°16’S, 166°55’E], VIII.1947, fr., J. Bernier 166 (P!) [+] ; c. 3 km NW of the Chute de la Madeleine, on Nouméa road, woodland in valley, growing with Agathis ovata and Gynostoma deplancheanum, on ultrabasic rocks, 22°12’S, 166°50’E, 17.III.1987, fl., K. L. Wilson 7024 (AD, MEL!, NSW!, NOU) [+] ; Creek Pernod, ravin au Sud du creek, Plaine des Lacs, sur serpentine, [22°11’30”S, 166°50’30”E], 16.XI.1950, imm. fr., fr., A. Guillaumin 8388 (G!, NY!, P!, RSA, Z[2 sheets]!) [+] ; Creek Pernod, Plaine des Lacs, route de Yaté, 150 m, [22°10’50”S, 166°50’34”E], 6.III.1966, fl. bud, fl., H. S. MacKee 14480 (CANB!, G!, K!, L!, MA, MO!, NOU!, NY!, P!, U!, Z!) [+] ; Creek Pernod, route de Yaté, maquis “carapace de fer”, 200 m, [22°10’50”S, 166°50’34”E], 23.II.1983, fl. bud, fl., H. S. MacKee 41264 (MO!, NOU!, P!) [+] ; Creek Pernod , 2.5 km N of creek, serpentine, [22°11’S, 166°51’E], 11.V.1977, fl., L. J. Musselman et al. 5099 (NOU!) [+] ; Creek Pernod, 2.3 km S of Route 2 bridge, [22°11’27”S, 166°50’30”E], 12.V.1977, fl., L. J. Musselman et al. 5129 (NOU!) [+] ; Creek Pernod , maquis, cuirasse ferrallitique, [22°10’50”S, 166°50’34”E], 4.III.2006, fl., Y. Pillon et al. 313 (MO!, NOU!) [+] ; Goro-Cascade , [22°17’30”S, 167°00’45”E], 20.IX.1978, imm. fr., B. Suprin 428 (NOU!) [+] ; Plaine des Lacs, NE of Grand Lac , low forest remnant, 300 m, [22°15’30”S, 166°55’30”E], 31.VIII.1980, imm. fr., G. McPherson 3008 (MO!, NOU!) [+ and *] ; Plaine des Lacs, E of Grand Lac , low forest remnants, [22°16’S, 166°56’E], 20.II.1983, fl. bud, fl., G. McPherson 5511 (MO!) [+ and *] ; Plaine des Lacs, E of Grand Lac , c. 3 km along road to Haut Kuebini , low forest, 300 m, [22°16’S, 166°55’E], 4.XI.1982, st., W. G. Ziarnik 49 (BRI!, NOU!) [+] ; route forestier de Mamié , maquis, [22°04’S, 166°54’E], 27.VII.1965, fl. bud, imm. fr., M. Schmid 498 (NOU!, P!) [□] ; Marais Kiki , colline entre les deux lacs, sur serpentine, 300 m, [22°09’30”S, 166°49’50”E], 26.IX.1950, st., M. G. Baumann-Bodenheim 6197 (K!, P!, Z!) [+] ; ibid., [22°11’S, 166°48’E], 26.IX.1950, fl., M. G. Baumann-Bodenheim 6231 (P!, Z!) [+] ; ibid., [22°09’30”S, 166°49’50”E], 26.IX.1950, fr., M. G. Baumann-Bodenheim 6251 (A!, NY!, P!, Z!) [+] ; Marais Kiki , route de Yaté, maquis haut, terrain serpentineux altéré, 200 m, [22°09’20”S, 166°49’50”E], 8.IX.1985, imm. fr., fr., H. S. MacKee 42795 (NOU!, P!) [+] ; Mont Kouakoué , maquis passant à fourré, 800 m, [21°57’S, 166°32’E], 18.XI.1972, fl. bud, M. Schmid 4291 (NOU!, P!) [□] ; Col de Ouénarou, route de Yaté, terrain serpentineux, maquis dégradé, 150 m, [22°10’S, 166°44’E], 19.III.1969, fl. bud, fl., H. S. MacKee 20293 (CANB!, K!, L!, MO!, NOU!, P!) [+] ; along Nouméa-Yaté road, c. 5 km E of Ouénarou (the turn-off towards the Rivière Bleue Reserve ), maquis vegetation, [160 m], [22°10’08”S, 166°44’35”E], 20.I.1980, fl. bud, fl., G. McPherson 2334 (MO!, NOU!) [+ and *] ; along Nouméa-Yaté road, c. 5 km W of Ouénarou , [22°10’S, 166°44’E], 21.III.1981, fl. bud, fl., G. McPherson 3609 (MO!, NOU!, P!, PTBG!) [+ and *] ; ibid., fl. bud, fl., G. McPherson 3610 (MO!, NOU!, P!, PTBG!) [+ and *] ; basse vallée de la Ouinne , 0-20 m, [21°59’S, 166°40’E], 3.VIII.1973, fl. bud, imm. fr., fr., H. S. MacKee 27132 (NOU!, P!) [□] ; Plaine des Lacs, fork of the road to Yaté, the left road leading back northward to 22 km station, [22°16’S, 166°55’E], 27.XI.1947, imm. fr., J. T. Buchholz 1430 (A!, NY!, US!) [+] ; Plaine des Lacs, route de Prony, [22°16’S, 166°55’E], 12.VIII.1977, imm. fr., T. Jaffré 1900 (NOU!, P!) [+] ; Plaine des Lacs, 200 m, [22°16’S, 166°55’E], 29.III.1912, fl., F. Sarasin 712 (Z!) [+] ; basse Pourina , forêt galerie, terrain serpentineux, 10 m, [22°01’S, 166°44’E], 7.XI.1979, fl. bud, H. S. MacKee 37554 (K!, L!, MO!, NOU!, P!) [□] ; Plaine des Lacs, haute Rivière Blanche , forêt des Électriques , forêt humide en vallée, terrain serpentineux, 200 m, [22°16’S, 166°55’E], 27.VII.1996, fr., H. S. MacKee 15365 (K!, L!, P!) [+] ; Rivière Blanche, galerie forestière en bordure de marais, [22°08’S, 166°40’E], 1965, fl., J.- M. Veillon 77 (NOU!, P!) [□] ; Rivière Bleue , maquis, 150 m, [22°06’S, 166°40’E], 1.VII.1965, imm. fr., L. Bernardi 9326 (G!, K!, L!, P!, Z!) [+] ; Haute Rivière Bleue , forêt avant le pont, sol alluvionnaire grossier, [22°06’S, 166°49’E], 4.IX.1963, fr., J. P. Blanchon 394 (NOU!, P!) [+] ; Parc Rivière Bleue , maquis near road just beyond where road crosses river (heading upstream), dense vegetation with trees and shrubs 3-8 m tall, porous-rocky, red soil, 230 m, [22°06’S, 166°39’E], 11.IV.1996, fl. bud, fl., J. C. Bradford & H. F. Hopkins 626 (MO!, NOU!, P!) [+] ; Rivière Bleue, maquis, 150 m, 1.VI.1994, fl. bud, J.- C. Pintaud 36 (P!) [+] ; près du pont de la route à Yaté sur la Rivière des Lacs , maquis serpentineux, [22°09’S, 166°51’E], 5.X.1950, imm. fr., A. Guillaumin & M. G. Baumann-Bodenheim 6483 (A!, G!, NY!, P!, Z!) [+] ; à l’ouest du pont de la route à Yaté sur la Rivière des Lacs , maquis serpentineux, [22°09’S, 166°51’E], 6.X.1950, fr., A. Guillaumin & M. G. Baumann-Bodenheim 6620 (L!, P!, RSA!, Z!) [+] ; Riviére des Lacs, 5 km en aval de la Chute , maquis, terrain serpentineux, 200 m, [22°09’S, 166°51’E], 1. III.1987, fl. bud, fl., H. S. MacKee 43461 (MO!, NOU!, P!) [+] ; Plaine des Lacs, serpentine slope, 6 km E of road parallelling Rivière des Lacs , 300 m, [22°13’S, 166°55’E], 2.VIII.1968, imm. fr., G. L. Webster & R. Hildreth 14479 (GH!, NOU!, P!) [+] ; Pont de la Rivière Madeleine ( Nouméa-Yaté ), maquis-fourré en bordure de la rivière, 150 m, 22°10’S, 166°50’E, 20.XII.1977, fr., P. Bamps 6015 (BM!, NOU!, P!) [+] ; sur la route à Yaté au PK 40, maquis serpentineux, 300 m, [22°09’S, 166°46’E], 21.II.1951, fl. bud, fl., A. Guillaumin & M. G. Baumann-Bodenheim 10747 (L!, Z!) [+] ; route de Yaté, 44 km from Nouméa, maquis, [22°10’S, 166°48’E], 5.XI.1959, fl. bud, R. F. Thorne 28584 (RSA!) [+] ; Yaté, [22°09’S, 166°54’E], 27.VII.1965, imm. fr., A. Aubr é ville 284 (P[2 sheets]!) [+] ; Yaté, Plateau au sud du village, maquis sur terrain serpentineux altéré, 300 m, [22°09’40”S, 166°54’40”E], 19.I.1975, fl. bud, fl., H. S. MacKee 29640 (BM!, CANB!, G!, K!, L!, MO!, P!, Z!) [+] ; Yaté [probably Lac de Yaté], 150 m, [22°10’S, 166°53’E], 20. IV.1987, fl., H. S. MacKee 43506 (NOU!, P!) [+] ; Yaté [probably village], [22°09’15”S, 166°54’40”E], 19.III.1912, fl., F. Sarasin 636 (Z!) [□]. — Province and Commune unassignable, “D”, [no date], st., A.- J. Le Rat 699 (P!) [□] ; “ Lre Western Track , maquis”, 240 m, 22.III.1982, fl., H. Brinon 1225 (NOU!) [□] ; “ Panlaithe ” or “ Panlaitche ” [E. Caldwell worked with E. Vieillard on the West Coast of New Caledonia in 1868, and Mrs. E. Caldwell sent his collections to K – fide Morat 2010], 1868, fl., E. Caldwell s.n. (K!) [□] ; [no specific locality], 27.X.1977, st., J. Bourret 1384 (NOU!) [□] ; [no specific locality], 186_ [final digit of year missing], fl. bud, E. Deplanche 284 [or 384?] (P!) [probable residual syntype of Solmsia calophylla ] [+] ; [no specific locality and no date], fl. bud, A.- J. Le Rat 1108 (P!) [□] ; [no specific locality, “ Donné par M. Pancher, 1870”], imm. fr., J. A. Pancher s.n. (P!) [probable residual syntype of Solmsia calophylla (uppermost branch), and possible residuals syntypes of Solmsia chrysophylla (two lower branches)] [+ and □] ; [no specific locality], 186_ [final digit of year missing], imm. fr., Petit 144 (P!) [possible residual syntype of Solmsia calophylla ] [□] ; [no specific locality and no date], st., Pennel 119 (MARS[2 sheets]!) [□] ; [no specific locality and no date], st., Pennel 428 (MARS!) [□].

DESCRIPTION

Shrubs (0.75-)1-4(-5) m tall, rarely treelets or small trees (0.5-)1-5(-6) m tall, dbh c. 10 cm (at 3 m) to 30 cm (at 5 m), branching dense or lax,± arcuate with a candelabra-like growth form; bark on older branches greyish or reddish brown, on dried material longitudinally striate and sometimes also transversely fissured/cracked; branches densely velutinous; pubescence tan-golden; trichomes very short; leaf scars on older stems conspicuous, subcircular, discolorous compared to surrounding stem.

Leaves

Alternate, usually only persisting near the distal tips of the branches, when internodes short sometimes giving a subopposite appearance, total leaf length (2.7-)3.5-11.5(-16) cm, smaller leaves usually associated with inflorescences, more proximal leaves generally larger, all leaves densely punctate; punctations usually translucent when fresh and blackish when dry, often obscured abaxially when densely velutinous (“chrysophylla form”); petioles (0.4-)0.7-2.5(-3.1) cm long, c. 1.5-2 mm in diam., longitudinally striate, robust, densely velutinous, pubescence initially tan-golden (darker brown on older petioles when dry), adaxially grooved, groove deeper near base of leaf blade; leaf blade obcordate or obovate, (2.2-)3-9(-13) × (0.8-)1.2- 5(-6) cm, blade length/width ratio c. 1.5-3.5(-4.5): 1, very thick, tough and fibrous, texture coriaceous; adaxial surface glabrous, very shiny, ± smooth or rugulose when fresh, more rugose when dry, darker green (or darker brown when dry) compared to abaxial surface; abaxial surface initially densely velutinous ( Fig. 6B ), remaining denser in “chrysophylla form” ( Fig. 7F ), usually becoming glabrescent in “calophylla form”; pubescence tan-golden or whitish; trichomes simple, very short, erect, soft; base cuneate or less often shortly attenuate-cuneate, more attenuated when margin more strongly revolute near base of blade; apex emarginate, retuse, or rounded; distal mucro c. 0.05(-0.1) mm long, relatively thick, formed by a short extension of the distal end of the midrib; margin usually slightly revolute, generally more obvious near base of leaf blade and after drying; midrib adaxially deeply grooved, abaxially very prominent and thick (c. 1.5-2 mm in diam. at midpoint of blade), longitudinally striate, densely velutinous; venation Calophyllum -like, c. 40-80(-100) vein pairs per side, adaxially generally invisible or usually only perceptible as rugulose folds, abaxially raised slightly and visible except when almost completely obscured by the dense velutinous pubescence (“chrysophylla form”), secondary veins diverging from midrib at an angle of (40-)60-70°, marginal nerve c. 0.05 mm from blade edge, vein course more arcuate in distal 1/4 of the blade; fine venation absent or inconspicuous.

Inflorescences

Similar in staminate and pistillate material ( Fig. 7D vs 7C, respectively); bracts absent or minute early caducous; peduncles 0.4-2.1 cm long; secondary inflorescence axes 1.5-11 mm long; floral clusters c. 5(-10)-flowered.

Flowers

Greenish-yellow, globose in bud, fragrant (× 5 reports on labels), densely velutinous, trichomes tan-golden or whitish; pedicels similar in staminate and pistillate flowers, longitudinally striate, densely velutinous, pubescence tan-golden, 1.8-2.5(-4.5) mm long (staminate pedicels), 1.5-2.5(-3) mm long (pistillate pedicels), and up to c. 3.5 mm long (fruiting pedicels); calyx similar in staminate and pistillate flowers, greenish-yellow; tube very short cupuliform or subcampanulate, c. 1-1.5 mm long; sepals subtriangular or ovate-triangular, c. 2 × 1.5-1.8 mm (staminate sepals), 2-2.5 × 1.5-2 mm (pistillate sepals), eventually reflexed or recurved in later anthesis, adaxially velutinous, often denser in distal half and especially near their somewhat thickened margins, densely velutinous abaxially, apex acute.

Staminate flowers. With (6)8(10) fertile stamens; filaments light green, (1.2-) 1.5-2 mm long, c. 0.2 mm wide, effects of compression and folding in S-shape in bud still visible during later anthesis ( Fig. 7B ); fertile anthers yellow-orange, c. 0.5- 0.6 × 0.5-0.6 mm (dry); pistillode minute, with all of the requisite parts but those greatly reduced in size.

Pistillate flowers. With (6)8(10) staminodia; staminodial filaments light green, 1-1.2(-1.4) mm long, c. 0.1-0.15 mm wide, distal portion generally recurved away from center of flower ( Fig. 7E ); sterile anthers punctate-subglobose, c. 0.1- 0.15 in diam.; ovary greenish, ± subglobose, (1-)1.5-1.8 × (1.2-) 1.5-2.2 mm, densely tomentose-sericeous, pubescence tan-golden or whitish; style 0.8-1 × 0.2-0.3 mm, glabrous, extending c. 0.05-0.1 mm beyond sepals; stigma discoid to subglobose, 0.5-0.6 × 0.7-0.9 mm, densely papillate, lobed ( Fig. 7E ).

Fruits

Greenish-yellow, obovoid or obpyramidal, 8-10 × 6.5-8.5 mm; base of undehisced fruit attenuated for c. 1-1.5 mm and subtended by the persistent (non-accrescent) calyx; apex of undehisced fruit depressed, slightly retuse to emarginate ( Fig.7C ); pericarp densely velutinous, pubescence tan-golden, persistent; persistent stamens, staminodia, style and stigma turning black (fresh or dry), remaining attached even after fruit valve dehiscence; dehisced fruit valves c. 6-8.5 × 5.5-6.5 mm.

Seeds

Black, c. 5.8-6.5 × 2 mm, covered with a thin transparent or translucent orange-yellow membrane ( Fig. 7F ), outer epidermis of membrane ± villous, trichomes orange-yellow, up to c. 1 mm long.

CONSERVATION ASSESSMENT

Solmsia calophylla is assigned a preliminary conservation assessment of Near Threatened, NT ( IUCN 2012). Using GeoCAT ( Bachman et al. 2011; http://geocat.kew.org/), and considering the aggregate populations of both morphogroups of S. calophylla , the Extent of Occurrence (EOO) is 14 008 km2 (Vulnerable, VU) and the Area of Occupancy is 296 km 2 (Endangered, EN), based on the recommended standard 2 × 2 km grid cell size ( IUCN 2019). The values of EOO and AOO qualify the species as VU using the primary Criteria B1 and B2, respectively, but the necessary subcriteria for B1 and/ or B2 are not met because S. calophylla is relatively widespread with plenty of suitable habitats, i.e., the habitat of the species is not severely fragmented, it occurs at many more sites than the 10 location maximum threshold, and there have been no extreme fluctuations detected in EOO, AOO, number of locations or subpopulations, or in the number of mature individuals (when analyzed in aggregate or as discrete morphogroups). Due to the preferences of S. calophylla to grow on ultramafics, many populations, occur within the “maquis minier”, a mining zone especially focused in the southern part of the island, and although maquis land is unsuitable for agriculture, it does occasionally become degraded by fires (Jaffré in ORSTOM 1981; Isnard et al. 2016). Populations of both morphogroups are well represented within New Caledonia’s protected area network, with plants already recorded inside at least 10 reserves and parks: Mont Do Special Fauna and Flora Reserve, Pic Ningua Special Botanical Reserve, Mont Humboldt Special Botanical Reserve, Mont Mou Special Botanical Reserve, Montagne des Sources Strict Nature Reserve, Parc Territorial de la Rivière Bleue, Les Lacs du Grand Sud Néo-Calédonien, Forêt Cachée Special Botanical Reserve, Pic du Pin Special Botanical Reserve, and Zone côtière Ouest Parc Provincial (UNEP-WCMC 2020). The preliminary conservation assessment of NT assigned to S. calophylla should be reevaluated periodically as stipulated by the IUCN, especially as nickel mining activities further encroach into suitable habitat of the species (see also L’Huillier et al. 2010).

Evaluating the conservation status of either informally recognized morphogroup (A-B) discretely proves problematic because of the ambiguous localities at Pouébo (for both morphogroups) and Canala (for Morphogroup A), which were each vouchered by a single herbarium collection made in the mid-1800s by Vieillard, a collector well-known for using confusing “irrational” labeling and numbering ( Morat 2010). Nevertheless, while it is true that analyzing Morphogroup A and B separately produce smaller values in important measures such as EOO and AOO for the two groups (especially for the EOO of Morphogroup A if the Pouébo locality is excluded from the distribution), the earlier unsatisfied subcriteria B1 and/or B2 mentioned before for the aggregate populations of the entire species still are not met for each morphogroup individually, and thus a level of threat higher than NT cannot be justified for either one.

TYPIFICATIONS

Baillon (1871) simultaneously published a composite genus description for Solmsia , along with detailed descriptions of its two constituent species, S. calophylla and S. chrysophylla , in a short article that appeared in volume 10 of his periodical Adansonia, Recueil d’Observations botaniques (dated 12 June 1871). The generic description included a mixture of comprehensive observations gathered from vegetative, flowering, and fruiting materials (including seeds). Domke (1934) effectively lectotypified S olmsia, by designating S. calophylla as its “Leitart.” The untypified status of the two binomials has never been addressed until the present study.

Baillon did not cite collection information for either species in the protologue, and only provided a very brief and similar general provenance for both. For Solmsia calophylla , the protologue locality was cited as “In sylvis montium Novae- Caledoniae”, the habit was noted as “arbor media”, and the flowers and fruits were described, whereas for S. chrysophylla , the locality was cited as “In montuosis Novae-Caledoniae”, the habit was given as “Arbor parva?”, and the inflorescences were generally described, without a specific description of the flowers or fruits.

In the introduction of the protologue article, Baillon (1871: 34, 37) specified that he obtained a fair amount of material to use in the genus and species descriptions from multiple sources via the statements “[...] de plusieurs explorateurs de la Nouvelle-Calédonie [...] La plupart de ceux auxquels nous avons eu recours nous ont remis [...] une autre plante qui paraît être bien commune dans le pays, car il n’y a pas de collection où elle n’abonde” [i.e., from several explorers who sent material of Solmsia along with specimens of another relatively widespread and closely related species endemic to New Caledonia — i.e., Lethedon tannensis ].

Particularly regarding original source material used for his second new species, Solmsia chrysophylla, Baillon (1871: 37) alluded to multiple collections gathered by those same unnamed botanical explorers who had provided him with material for the protologue description of S. calophylla , via the passage “Dans une autre Espèce [= S. chrysophylla ], également abondante dans les collections, [...]” [i.e., in the other species (= S. chrysophylla ), equally abundant in the collections [...]].

Determining the identities of Baillon’s “de plusieurs explorateurs de la Nouvelle-Calédonie ” and identifying their collections that served as original material for the Solmsia species is simplified by the fact that only a few botanists, mostly French in origin, had collected plant specimens from New Caledonia by mid-1871 ( Guillaumin 1911; Morat 2010), and even fewer still visited the regions of the Grande Terre where Solmsia occurs.

Considering published itinerary information taken together with the material deposited in the P herbarium, Baillon’s home herbarium, there were four significant botanists who collected potential syntype specimens: Eugène Vieillard (coll. 1855-1867, used “numérotation spéciale et irrationnelle” [= special and irrational specimen numbering]), Jean Armand Pancher (coll. 1857-1869, also used special and irrational specimen numbering), Émile Deplanche (coll. 1855-1860 and 1861-1867, used “particulière” [i.e., peculiar/irregular] specimen numbering), and Benjamin Balansa (coll. 1868- 1872) (parenthetical information fide Morat 2010). It is interesting historically to note that material of Solmsia , a relatively common and widespread plant, was not collected by the 18th century explorers, including the first Europeans to visit New Caledonia, i.e., The Forsters and W. Anderson (coll. 1774), who accompanied Captain James Cook on his second voyage on the H. M. S. Resolution , and J. Labillardière (coll. 1793), another prolific early visitor who led the second expedition to New Caledonia as part of a mission to find out what happened to the lost French expedition led by Lapérouse.

The confusing and often inconsistent labeling and specimen numbering systems present on Vieillard, Pancher and Deplanche duplicates, widely distributed to many herbaria around the world, are problematic on the examined potential syntype sheets of Solmsia at P. For instance, individual Vieillard sheets often contain multiple specimens belonging to both morphogroups, while some identically numbered sheets may bear more than one label (or specimen) that frequently includes at least two contradictory localities (e.g., “Kanala” vs “Kanala, Mt. Dore” vs “Kanala, etc.” vs “Mont-Dore” vs “Mont-Dore[deleted anonymously, and replaced with] Kanala”), or dates (e.g., labels typewritten as “1855-1860” vs “1861-1867”, and sometimes with a second handwritten year that falls outside of the year range printed on the typewritten label). One specific worst-case example of Solmsia labels with these kinds of discrepancies besides other irregularities in collection numbering is observable in the Vieillard “duplicates” with numbers variously handwritten as “ 145, 146 ”, “ 145=146 ”, or simply either “ 145 ” or “ 146 ” that may be present on one or more labels attached to an individual sheet. In some instances, a sheet with a single number may have two or more affixed specimens belonging to the two different morphogroups, or conversely a sheet numbered with two different numbers, e.g., containing both “ 145 ” and “ 146 ”, may only bear a single affixed branch. Regarding the examined Pancher material, one unnumbered sheet (P06622220!) was nicely annotated by Baillon as S. calophylla and with its partial place of publication (as “ Solmsia calophylla H. Bn in Adans. X. 38.”), but only the uppermost branch of the three branches affixed to this particular sheet corresponds taxonomically to his S. calophylla . In fact, it is certainly possible that the two lowermost branches on P06622220 may actually represent syntype material of Baillon’s S. chrysophylla , which coincidentally also first appeared on the same protologue page ( Baillon 1871: 38) as S. calophylla . It is not clear if Baillon missed that the specimens were mixed from different collections, or if the mistake was introduced after the protologues were published, perhaps during the mounting process. Regardless, the dubiously determined, mixed, mostly sterile nature of these Pancher specimens make them undesirable choices as lectotypes for either binomial.

The fourth New Caledonian explorer, Benjamin Balansa, collected several Solmsia collections that do not suffer from the kinds of problems plaguing the specimens of the three other botanists. Balansa assigned handwritten unique collection numbers to his material following a standard numerical format. In addition, Balansa’s collections are usually larger, more fertile, and contain more detailed label information, such as descriptions of habitat and habit, and clearly specified collection dates. Given these advantages, two Balansa collections at P, coincidentally with adjacent collection numbers, 263 and 262, are the most suitable choices for the lectotype collections of Baillon’s S. calophylla and S. chrysophylla , respectively. Moreover, both collections bear almost identical labels handwritten by Balansa citing the exact same locality (“Collines ferrugineuses de la Baie du Prony”), habit/height description (“Arbrisseau de 2 mètres de hauteur”), and date (“ Septembre 1868 ”), which again illustrate the similarities related to habit, ecology and distribution found in Morphogroups A and B.

Specifically regarding the lectotypification of Solmsia calophylla , both P sheets of Balansa 263 (P00239783!, P06622659!), each bear a single, large nice quality fruiting branch that closely matches the morphology described in the protologue ( Baillon 1871). Sheet P00239783 ( Fig. 8 ) is designated as the lectotype because it is the only duplicate of the two that includes the species annotation in Baillon’s handwriting (as “ Solmsia calophylla H. Bn ”, but without date of determination and protologue publication details). Two labels are affixed to the sheet. The first label includes Balansa’s handwritten inscription: “ 263. Arbrisseau de 2 mètres de hauteur. Collines ferrugineuses de la Baie du Prony (Nlle Calédonie). Septembre 1868 ”, whereas the second label was typewritten as “HERB. MUS. PARIS. NOUVELLE CALÉDONIE. M.BALANSA, 1868-1870.” Baillon did not mention some of Balansa’s specific label details in his protologue, possibly because Baillon had examined so much material belonging to his new species, or perhaps he may have been aware of numbering/labeling problems so prevalent in Vieillard material.A third possibility is that he primarily studied the ample fruiting specimen affixed to the sheet designated here as an isolectotype (P06622659), which bears minimal inscriptions on its two labels (viz a handwritten collection number by Balansa on one label, and an anonymous handwritten mention of Nouvelle-Calédonie on the second label).

For Solmsia chrysophylla , all three sheets (P00239812!, P06622240!, P06622241!) of the designated lectotype collection, Balansa 262, have large, nice quality, fertile specimens, complete with many mature, dehisced fruits and floral buds, and with all branches closely matching the morphology described in the protologue ( Baillon 1871). Sheet P06622241 ( Fig. 9 ) is specifically chosen as the lectotype because it is the only one of the three duplicates to include the species annotation in Baillon’s handwriting (as “ Solmsia chrysophylla H. Bn ”, but without date of determination and protologue publication details). Sheet P00239812 is the only one of the three sheets to include specific information relating to locality, plant height, habitat, and date of collection, via a label with Balansa’s handwritten inscription: “ 262. Arbrisseau de 2 mètres de hauteur.Collines ferrugineuses de la Baie du Prony. Septembre 1868. B. Balansa.” The second label is the same kind of “HERB. MUS. PARIS” typewritten label that was affixed to the lectotype designated for S. calophylla (Balansa 263). Finally, isolectotype sheet P06622240 bears minimal inscriptions on its two labels, most notably Balansa’s handwritten collection number “ 262 ”, but it also contains a single large fruiting branch affixed to the sheet that is undoubtedly duplicate material of the lectotype sheet.

Counting the lectotypes and isolectotypes of Solmsia calophylla and S. chrysophylla , a total of c. 55 sheets (representing c. 15 different collections) were identified as either probable or possible syntype material of Baillon’s two binomials (c. 30% of the sheets are deposited in other herbaria besides P and those are thus regarded as isosyntypes). All residual syntype information is indicated inside square brackets immediately after the relevant repository information cited in the Specimens examined section (exsiccatae details including barcode/accession numbers and the images of the syntypes are posted at https:// www.tropicos.org/ and https://science.mnhn.fr/all/search).

Figures

FIG. 1. — Solmsia Baill.: A, branch from pistillate plant, with immature fruits and unopened floral buds; B, staminate floral bud, showing S-shaped, folded filaments; C, D, stamen; E, pistillate flower; F, sepals and several associated staminodia removed from a pistillate flower; G, gynoecium; H, ovary, longitudinal section; I, very immature loculicidal capsule; J, immature seed. Modified from plate of figures originally published in Schlechter (1906). All illustrated parts based upon a mixture of specimens gathered from staminate and pistillate plants collected under F. R. R. Schlechter 15138 (BM!, G!, K!, L!, P!, Z!). Scale bars:A, 6 mm; B, 1 mm; C, D, 0.5 mm; E-H, 1 mm; I, 3 mm; J, 2 mm.

FIG. 6. — Solmsia calophylla Baill. (photos A-D belong to plants of Morphogroup A,“calophylla form”; photo E belongs to a plant of Morphogroup B, “chrysophylla form”): A, B, habit and vegetative features at La Madeleine (photos: B. Henry, 23.VIII.2009); C, habit at Forêt Cachée (photo: B. Henry, 5.II.2012); D, branches with immature fruits (photo: B. Suprin, locality and date unavailable); E, branch, Barrage de Dumbéa (photo: C. Davidson, 11.XI.2007, based on Munzinger et al. 4666, NOU!, P!).Photos A-D downloaded from https://endemia.nc/, copyright of the photographers,and covered under Creative Commons Attribution-Non Commercial 4.0 International Public License (CC BY-NC 4.0), viewable at: https://creativecommons.org/licenses/by-nc/4.0/legalcode [accessed 24.X.2020]. Photo E kindly provided with compliments of the photographer.

FIG. 7. — Solmsia calophylla Baill. (photos A, B belong to plants of Morphogroup A, “calophylla form”; photos C, D, F belong to plants of Morphogroup B, “chrysophylla form”; photo E lacks vegetative material and locality details and thus cannot be placed in either morphogroup): A, B, branches with inflorescences of a staminate plant at Forêt Cachée (photos: B. Henry, 5.II.2012); C, immature loculicidal capsules at Dumbéa Nord (photo: G. Gâteblé, 1.IX.2005); D, flowering branch from a staminate plant at Col de Plum (photo: G. Gâteblé, 15.II.2006); E, flowers of pistillate plant with an ant visitor (photo: B. Suprin, locality and date unavailable); F, branch with one mature, 4-carpellate capsule (with three seeds visible) at Barrage de Dumbéa (photo: C. Davidson, 11.XI.2007, based on Munzinger et al. 4666, NOU!, P!). Photos A-E downloaded from https://endemia.nc/, copyright of the photographer, and covered under Creative Commons Attribution-NonCommercial 4.0 International Public License (CC BY-NC 4.0), viewable at: https://creativecommons.org/licenses/by-nc/4.0/legalcode [accessed 24.X.2020]. Photo F kindly provided with compliments of the photographer.

FIG. 8. — Lectotype of Solmsia calophylla Baill. designated in this study (B. Balansa 263, P00239783!). Scale bar: 4 cm.

FIG. 9. — Lectotype of Solmsia chrysophylla Baill. [= S. calophylla Baill.] designated in this study (B. Balansa 262, P06622241!). Scale bar: 4 cm.

FIG. 2. — Distribution of Solmsia Baill. in New Caledonia. Morphogroups: A, “calophylla form” (+); B, “chrysophylla form” (Oi). The two arrows indicate doubtful sympatric populations of both morphogroups based on questionably labeled Vieillard material collected in the mid-1800s: upper arrow denotes the Pouébo locality (non-ultramafic substrate) where both morphogroups were likely erroneously reported on the label of E. Vieillard 254 (P00239877!); lower arrow denotes the “Kanala” [= Canala] locality where several ambiguously labeled sheets of Morphogroup A were supposedly collected (e.g., Vieillard 145, “145=146” and “147”). See Results and discussion section for more details.