Adelobotrys microcarpus Schulman, 2015

Schulman, Leif & Ruokolainen, Kalle, 2015, Adelobotrys tessmannii (Merianieae, Melastomataceae) and allies: a refined circumscription and description of two new Amazonian species with notes on their ecology, Phytotaxa 234 (2), pp. 101-120 : 113-117

Adelobotrys microcarpus Schulman , sp. nov. ( Fig. 6 )

Type: — PERU. Loreto: Alto Amazonas, High rainforest along Río Marañon near Teniente Pinglo , just above Pongo de Manseriche , 250–300 m, 4–7 October 1962 (fl., fr.), J.J. Wurdack 2070 (holotype NY!; isotypes F!, G!, K!, S!, US!).

Diagnosis: — Adelobotrys tessmannii Markgr. affinis sed laminis fere glabris marginibus irregulariter serrulatociliolatis, partibus ultimis inflorescentiae cincinnis duplicatis ramis perelongatis, floribus minoribus, dentibus exterioribus calycis obsoletis, et hypanthiis in fructu parvis globosis differt.

Allied to Adelobotrys tessmannii Markgr. but differs by having almost glabrous laminae, lamina margins irregularly serrulate-ciliolate, ultimate branching of the inflorescence in the form of double cincinni with strongly elongated branches, smaller flowers, obsolete outer calyx teeth, and small globose fruiting hypanthia.

Full description: — Liana adhering to support by adventitious roots to ca. 5 cm long produced in comb-like rows on internodes, reported height of fertile plants (1–)3–10(–15) m. Stem slender, diameter in fertile branches to 0.5 cm, terete to somewhat fluted, when very young densely pubescent with dull- to dark-brown, appressed, short (ca. 0.4–1 mm long) malpighian hairs, soon largely glabrescent; stem surface between hairs smooth, dull straw-coloured to brown or greyish; internodes in vegetative parts of fertile stems 4–10 cm long; nodes hardly swollen, not more pubescent than the internodes (apart from densely pubescent axillary buds). Leaves opposite, isomorphic. Petiole (1.2–)2.2–4.8(–6.2) cm long (ratio of lamina length to petiole length ranges from 2.3 to 7.5 but is typically around 4.5), ca. 1 mm in diameter; terete but flattened to somewhat grooved above; sparsely to fairly densely covered with brown malpighian hairs at first, glabrescent with age especially below. Lamina narrowly ovate to elliptic, (7.5–)8.5–14.5(–16) × (4–)5–7(–8) cm (ratio of length to width 1.7–2.3), base rounded to slightly cordulate, apex acuminate (to almost acute), margin mostly serrulate, often somewhat revolute (in herbarium specimens); chartaceous; in herbarium specimens typically greyish-olive-green with darkish brown veins below, somewhat shiny medium green in living plant. Both surfaces of lamina pubescent with medium- to dark-brown malpighian hairs at first but very soon glabrescent, mature leaves glabrous above (or with a few hairs along main veins), below sparse to medium pubescence remains along main and transverse veins and scattered hairs between them, the hairs being appressed, 0.5–1 mm long, laterally flattened but not dilated (apparently 1 cell layer thick, 2 layers wide), symmetrical to somewhat asymmetrical, sessile to very short-stalked and substraight to slightly sigmoid especially on main vein (and then broadly V- rather than T-shaped); on lamina proper on abaxial side malpighian hairs interspersed with minute, elongate-club-shaped glands (ca. 0.05–0.1 mm long); epidermis of adaxial surface usually with rather dense greyish hemispherical protuberances ca. 0.07 mm across. Margin of lamina with sparse ciliolation of tangential malpighian hairs and with persistent, obliquely spreading, pale, multiseriate, terete, unbranched, 0.5–0.7 mm long setulae tipping marginal teeth 1–2 mm apart. Venation basally acrodromous with midrib plus four distal main veins, stronger pair of acrodromal veins only somewhat closer to lamina margin than to midrib (distances 1:1–1:1.8), pinnate secondary veins visible below, 0.5–0.8 cm apart; tertiary veins not visible; veins prominent below, flat to somewhat impressed above. Axils of main veins on abaxial side with a few multiseriate, terete, unbranched setulae 0.5–1 mm long, tipped by a multicellular gland, a few such setulae sometimes also present on midrib a couple of cm above the base. Inflorescence a terminal, broadly conical, lax double thyrsoid with up to hundreds of flowers; at anthesis (16–)22–25 × 12–18 cm including (4–) 6–7.5 cm long peduncle; rachis with 4 elongated internodes, nodes with two main branches consisting of usually 2 or 3 elongated internodes and with two shorter branches (of 1–2 internodes) produced from accessory buds, internodes of rachis, and branches, progressively shorter towards apex; partial inflorescence a double cincinnus with branches bearing up to 12 flowers each, characteristically elongated in fruit and reminiscent of arms of an octopus when only scars of fallen fruits remain. Peduncle subtended by leaves similar to ordinary vegetative leaves; lowermost node of rachis with bracts similar to but smaller than vegetative leaves, upper nodes with progressively smaller and narrower (sub)sessile bracts, bracts at lower nodes persistent but those at upper nodes mostly absent already at anthesis; cincinni subtended by caducous sessile elliptic bracteoles 3–5 × 2–3 mm. Pedicel at anthesis 3–5 mm, slender. Flower reportedly with slight sweet fragrance, perigynous, hypanthium coriaceous, cylindrical to ellipsoid, smooth, rather abruptly widening from pedicel, 2.5–3 mm long and 1.5–2 mm wide at torus; calyx spreading-cup-like, shallowly 5-lobed, ca. 1.6 mm long; outer calyx teeth absent to visible as vestigial knobs (to ca. 0.4 mm long), asetose; pedicel with dense, hypanthium and calyx with sparse cover of brown malpighian hairs, hypanthium glabrescent in fruit, calyx sparsely pubescent also on inner side but with rim mostly aciliate; hypanthium reportedly green in living plant but calyx light orange-brown to even purple or blue. Petals 5, colour in living plant reportedly (whitish to) pink or purple to lilac or even blue, when dry dull yellow, glabrous; obovate, 5.9–7.1 mm long; venation actinodromous and flabellate. Stamens 10, anisomorphic, bent and crowded to one side of flower, glabrous; filaments ca. 4 mm, laterally dilated, strap-like, ca. 0.3 mm wide; geniculate at junction between filament and anther, anther bent introrsely (i.e., with ventral side down). Anthers reportedly yellow, subulate, dorsal-arcuate, base rounded; single apical pore ca. 0.2 mm across; connective hardly prolonged between filament and thecae, basally extended into an erect triangular spur, dorsally into a gradually ascending, apically deeply cleft appendage, which is reportedly rosy pink. Larger (epipetalous) stamens: thecae clearly arcuate (sometimes even J-shaped), 4.9–5.5 mm long and 0.4 mm thick at base, apical pore dorsally inclined; basal connective appendage ca. 0.6–0.7 mm long, subulate and variously slightly lobulate, dorsal appendage 2.2–3.2 mm long, cleft by ca. 0.5– 0.8 mm. Smaller (episepalous) stamens: thecae only slightly arcuate, 3.1–3.9 mm long and 0.4 mm thick at base, apical pore subterminal; basal connective appendage ca. 0.5 mm long, irregularly lobulate, dorsal appendage 2.2–2.6 mm long, cleft by ca. 0.5–0.7 mm. Ovary 5-locular, roughly cylindrical, somewhat fluted, truncate at apex, 2.6–3 × 1.2–1.5 mm, glabrous; style 4.9–5.9 mm long, somewhat sigmoid, glabrous; stigma punctate. Pedicel in fruit to 5–7 mm. Fruiting hypanthium globose, clearly constricted at torus, 2.5–4.5 × 3–4.5 mm; thin, sparsely pubescent, 10-costate, disintegrating at maturity revealing costae that are not apically connected by a circular vascular strand. Calyx persistent in fruit, horizontally spreading, as wide as hypanthium, 1.4–1.6 mm long. Fruit a 5-locular loculicidal capsule completely enveloped by the hypanthium but free from it; globose, somewhat 5-angled, rounded-truncate at apex. Placenta central, sessile and hardly extended from the central column. Seeds numerous, narrowly wedge-shaped, ca. 0.9–1 × 0.25 mm; surface smooth to slightly rugose; embryo occupying central third, testa elongated at both ends possibly forming air-filled sacs; testa light brown, embryo darkish brown (in herbarium specimen).

Notes: —This species was marked as Adelobotrys tessmannii morphotype 2 in the phylogenetic analysis by Schulman & Hyvönen (2003).

Habitat: —Primary to somewhat disturbed or secondary lowland (to premontane) rainforest, mostly in undulating terrain on lateritic loamy soil. Elevation from 100 to 800 m a.s.l.

Distribution: —Western Amazonia between 67°47’ and 78°40’ W, from 3°18’ S in Loreto to 12°21’ S in Madre de Díos, Peru, and in western Brazil ( Fig. 3 ).

Etymology: —The epithet was chosen because this species has notably small fruits, smaller than any other known species in the genus.

Phenology: —Collected with flowers in January–October; with fruits in January–March and May–October.

Conservation: —Least concern (LC), i.e., not threatened. The extent of occurrence clearly exceeds the threshold value of 20,000 km 2. The available data are too scanty to accurately estimate the true area of occupancy and, especially, population size or trends in it, but there is no plausible reason to estimate that these would approach levels warranting placement in or near threatened categories. However, of the three species treated here, A. microcarpus has the smallest extent of occurrence and is, according to currently available data, near-endemic to Peru.

Paratypes: — BRAZIL. Amazonas: ca. 2 km north from the village of Lago do Pupunha, Río Juruá , 100 m, 5°34’ S 67°47’ W, 1 April 2012, Ruokolainen, K. with H. Tuomisto, T. Emilio, F. Figueiredo & G. Moulatlet 15966 ( TUR!). PERU. Amazonas: Bagua, Imaza, Camino de Yamayakat hacia Kuzú, 270 m, 18 July 1994, Díaz, C., Jujúa Katip, S. & Peña, A. 6923 ( MO!) ; Amazonas: Bagua, Imaza, Región Nororiental del Marañon , Comunidad de Kampaenza , Ribera de la quebrada Shimutaz. , Río Marañon , 320 m, 4°55’ S 78°19’ W, 7 October 1994, Jaramillo, N. & Peas, C. 533 ( MO!) ; Amazonas: Bagua, Imaza, Región del Marañon, Comunidad de Yamayakat, Río Marañon , 300 m, 4°55’ S 78°19’ W, 20 January 1995, Rodríguez, E. 210 ( MO!) ; Amazonas: Bagua, Imaza, Comunidad de Kampaensa , arriba río Shimutaz , 450 m, 4°55’ S 78°19’ W, 22 October 1995, Rodríguez, E. 611 ( MO!) ; Amazonas: Bagua, Imaza, Yamayakat , 350 m, 5°3’20” S 78°20’23” W, 1 June 1996, Vásquez, R. & Vásquez, A. 21032 ( MO!) ; Amazonas: Bagua, Imaza, Kampaensa , 320 m, 4°55’ S 78°19’ W, 23 October 1995, Vásquez, R., Jaramillo, N. & Mayan, C. 20406 ( TUR!) ; Amazonas: Bagua, Senepa, 2nd road from Nuevo Nazareth ( Imacita ), near mouth of Río Imaza , on Río Marañon , leading inland 2 km to Carretera , 250–300 m, 26 January 1967, Tillett 671-24 ( US!) ; Amazonas: Bagua, Yamayakat , trocha a Putuim , 400 m, 4°55’ S 78°19’ W, 16 October 1996, Vásquez, R. & Jaramillo, N. 20287 ( MO!) ; Amazonas: Condorcanqui, El Cenepa , Comunidad de Mamayaque , 400 m, 4°34’49” S 78°14’1” W, 9 August 1997, Rojas, R., Peña, A. & Chávez , E. 0251 ( MO!) ; Amazonas: Condorcanqui, El Cenepa , Comunidad de Mamayaque , Cabecera de la quebrada Saasá , 560 m, 4°37’3” S 78°19’57” W, 13 August 1997, Vásquez, R. & Quiaco, E. 24548 ( MO!) ; Amazonas: Condorcanqui, El Cenepa , Comunidad de Tutino , 340 m, 4°33’5” S 78°12’54” W, 28 July 1997, Vásquez, R., Ampam, D., Quiaco, E., Ampam, A. & Dupis, C. 24450 ( TUR!) ; Amazonas: Condorcanqui, El Cenepa , Comunidad de San Antonio, Río Cenepa , 300 m, 4°29’30” S 78°10’30” W, 20 June 1997, Vásquez, R., Peña, A., Chávez, E. & Quiaco, E. 24078 ( TUR!) ; Amazonas: Condorcanqui, El Cenepa , Comunidad de Mamayaque, Río Cenepa , Quebrada Sása , 400 m, 4°37’8” S 78°13’46” W, 6 February 1997, Vásquez, R., Rojas, R. & Peña, A. 22385 ( MO!) ; Amazonas: Condorcanqui, El Cenepa , Comunidad de Mamayaque, Río Cenepa , Quebrada Sása , 300 m, 4°31’40” S 78°11’40” W, 26 January 1997, Vásquez, R., Rojas, R., Peña, A., Chávez, E. & Quiaco, E. 22337 ( MO!) ; Amazonas: Río Cenepa, vicinity of Huampami , ca. 5 km E of Chávez Valdívia, el camino de Etseketa-Río Cenepa , 200–250 m, 4°30’ S 78°30’ W, 10 August 1978, Ancuash, E. 1381 ( MO!) ; Amazonas: Río Cenepa, vicinity of Huampami , ca. 5 km E of Chávez Valdívia, alrededor de la comunidad Huampami , 200–250 m, 4°30’ S 78°30’ W, 12 August 1978, Ancuash, E. 1439 ( MO!) ; Amazonas: Quebrada Kayamas lugar Cenepa , 390 m, 6 April 1973, Ancuash, E. 186 (HUH!, MO!) ; Amazonas: Quebrada Huampami , 213 m, 5 May 1973, Ancuash, E. 328 (F!, HUH!, MO!) ; Amazonas: Quebrada chigkan entsa, Río Cenepa , 390 m, 4°30’ S 78°30’ W, 9 June 1973, Ancuash, E. 590 (F!, HUH!, MO!) ; Amazonas: S of Huampami trail to house of Theodora, S of Río Cenepa , 240–255 m, 17 July 1974, Berlin, B. 1669 (HUH!, MO!) ; Amazonas: Río Cenepa ; 10 km So. of Huampami , 300 m, 16 September 1972, Berlin, B. 53 (HUH!, MO!) ; Amazonas: Monte al lado de Huampami , 240–255 m, 29 July 1974, Kayap, R. 1338 (HUH!, MO!) ; Amazonas: Río Cenepa, vicinity of Huampami , ca. 5 km E of Chávez Valdívia, al lado de casa de Vicente , Quebrada Apigkan entsa, 200–250 m, 4°30’ S 78°30’ W, 3 August 1979, Kujikat, A. 128 (F!, MO!) ; Amazonas: Río Cenepa, vicinity of Huampami , ca. 5 km E of Chávez Valdívia, alrededor de casa Comunidad , cerca de Huampami , 200–250 m, 4°30’ S 78°30’ W, 12 August 1978, Kujikat, A. 300 (F!, MO!) ; Amazonas: Río Cenepa, vicinity of Huampami , ca. 5 km E of Chávez Valdívia, al lado de Chigkan entsa, Quebrada Aintami , 200–250 m, 4°30’ S 78°30’ W, 17 August 1978, Kujikat, A. 410 ( MO!) ; Huánuco: Pachitea, Codo de Pozuzo , alluvial fan floodplain of Río Pozuzo after it emerges from mountains, trail to NW behind settlement, 450 m, 9°40’ S 75°25’ W, 18 October 1982, Foster 9258 (AAU!, TEX!, MO) ; Huánuco : Pachitea ; Codo de Pozuzo, alluvial fan floodplain of Río Pozuzo after it emerges from mountains, trail N of settlement to Río Mashoca , 500 m, 9°37’ S 75°25’ W, 19 October 1982, Foster 9293 (NY!, TEX!, US!, MO) ; Loreto: Loreto, Nauta , from km 9.5 of Carretera Nauta-Iquitos ca. 1 km E-wards, 100–200 m, 4°26’44” S 73°35’5” W, 22 September 1998, Schulman, L. 212 (AMAZ!, TUR!, USM!) ; Loreto: Mariscal Ramon Castilla, ca. 2 km SW from the village of Puerto Izango , río Yaguasyacu , terrace of río Amazon , 100–150 m, 3°18’ S 72°1’ W, 19 May 1997, Ruokolainen, K., Tuomisto, H., Vargas, V. & Vormisto, J. 11122 ( TUR!) ; Loreto: Maynas, ca. 3 km SE from the village of Sta Ana , 150–200 m, 4°6’ S 73°7’ W, 3 February 1996, Ruokolainen, K., Criollo, G., Sarmiento, A., Tuomisto, H., Koivunen, H., Lusa, E. & Vormisto, J. 8927 ( TUR!) ; Loreto: Maynas, River Momón , near to the tourist lodge of Amazon Selva Tours , 100–150 m, 3°40’ S 73°20’ W, 19 May 1986, Ruokolainen, K., Tuomisto, H., García, M., Ríos, R. & Torres, A. 417 ( TUR!) ; Loreto: Balsapuerto ( lower Río Huallaga basin), 350–550 m, 29 August 1929, Killip, E. P. & Smith, A. C. 28501 (NY!, US!) ; Loreto: Balsapuerto ( lower Río Huallaga basin), 150–350 m, 28–30 August 1929, Killip, E. P. & Smith, A. C. 28634 (HUH!, NY!, US!) ; Loreto: Balsapuerto , 220 m, March 1933, Klug, G. 2948 (BM!, F!, G!, HUH!, K!, NY!, S!) ; Madre de Dios: Manu, Close to the village of Diamante , southern side of Río Alto Madre de Dios , 400 m, 12°21’ S 70°57’ W, 23 October 1998, Ruokolainen, K., Tuomisto, H. & Zegarra, A. 13412 ( TUR!) ; San Martín: San Martín, km 28 of Tarapoto-Yurimaguas road, 750–800 m, 6°27’ S 76°19’ W, 23 September 1986, Knapp, S. & Mallet, J. 8386 ( MO!) ; San Martín: Pongo de Cainarachi, Río Cainarachi , tributary of Río Huallaga , 230 m, Sep.–Oct. 1932, Klug, G. 2603 (BM!, F!, G!, HUH!, K!, NY!, S!, US!) ; Eastern Peru, Tessmann 3951 (F!, NY!) ; Stromgebiet des Maranon von Iquitos aufwärts bis zur Santiago-Mündung am Pongo de Manseriche , 4°28’0” S 77°37’0” W, 1924, Tessmann 4076 (G!, S!) .

Specimens of uncertain status determined as A. cf. microcarpus : — BRAZIL. Amazonas: ca. 3 km north-west from the village of Roque, Río Juruá , 100 m, 5°5’ S 67°14’ W, March 28 2012, Ruokolainen, K. with H. Tuomisto, T. Emilio, F. Figueiredo & G. Moulatlet 15942 ( TUR!) ; Amazonas: Manaus-Caracaraí road, km 72, area of WWF project “Minimal critical size of Ecosystems”, near streamlet of the camp of area “1302”, 30 April 1984, Renner 914 ( AAU!) . PERU. Loreto: Requena, ca. 10 km E from Jenaro Herrera , 150–200 m, 4°55’ S 73°35’ W, 18 September 1998, Ruokolainen, K., Tuomisto, H. & Cardenas, G. 12914 ( TUR!) ; Loreto: Requena, ca. 2 km N from the biological station of Jenaro Herrera , 150–200 m, 4°52’ S 73°39’ W, 18 September 1998, Ruokolainen, K., Tuomisto, H. & Cardenas, G. 12897 ( TUR!).—all are sterile, young specimens with insufficient diagnostic characters for unequivocal identification .

Soil preferences: —Of the species treated here, A. latifolius was the most frequently observed, with 54 occurrences in a total of 396 inventory transects in the four geographical areas ( Fig. 1 ). The frequency of A. microcarpus was less than one third of that (14 transects), and the least frequent was A. tessmannii (6 transects). Although all three species are broadly sympatric in the sense that their general distribution areas overlap, they tend not to share the same habitats on a local scale. Adelobotrys microcarpus was never observed to co-occur with the two other species, and A. latifolius and A. tessmannii co-occurred in only one transect.

Adelobotrys latifolius was typically found in forests on soils with relatively high cation concentration, whereas A. microcarpus tended to occupy forests on poorer soil ( Fig. 7 ). This pattern was found both in regions where the two species co-occurred (Loreto and Juruá) and in the regions where only one of the two species was found (Madre de Dios and Yasuní). The probability to obtain in Loreto and Juruá by chance an equally or more strongly different average soil cation value for two species with the frequencies observed for A. latifolius and A. microcarpus were 0.022 and 0.314, respectively. The random probability that these minimum differences coincide and show the same direction (the same species is occupying poorer soils in both regions) is (0.022 × 0.314)/2 ≈ 0.003. Accordingly, we can refute the null hypothesis that the species would have no difference in their preference for soil cation concentration. This conclusion is reinforced by the observed occurrence patterns from regions where the two species were found alone. No statistically significant difference in preferred soil cation concentration was found between A. tessmannii and A. latifolius (p = 0.304), or A. tessmannii and A. microcarpus (p = 0.318).

Most of the inventory transects were established in unflooded (terra firme) forests, but 35 of them (12 in Madre de Dios, 10 in Loreto and 13 in Yasuní) were established in floodplain forests that were either well-drained or swampy and dominated by the palm Mauritia flexuosa . Adelobotrys microcarpus and A. tessmannii were exclusively found in terra firme, while A. latifolius was found in seven floodplain transects in Yasuní, even though in Loreto also this species was encountered only in terra firme transects.

We have not tried to analyze possible preferences of the species in relation to topographic variation within the inventory transects. However, A. latifolius was clearly found more often along small creeks compared to the other two species.

Figures

FIGURE 1. Localities (n=390) of quantitative species inventories (n=396) and soil sampling for estimating soil preferences of the species treated here. Individuals of the species were found in the localities as follows: orange dot = Adelobotrys latifolius, green dot = A. microcarpus, violet dot = A. tessmannii; in one locality two species were found. The four separate regions in which field studies were carried out and where at least one of the three species was encountered are marked (Madre de Dios in southern Peru, Loreto in northern Peru, Yasuní in Ecuador and Juruá in central Brazil). Elevation is depicted as follows: 0–1000 m as white; 1001–2000 m, 2001–3000 m, 3001–4000 m, and>4000 m as progressively darker shades of grey. For approximate location of map section in South America, see Fig. 3D.

FIGURE 3. Collecting localities of specimens of the species treated here. A. Adelobotrys tessmannii (solid triangles), A. cf. tessmannii (open triangles), and possible hybrids between A. tessmannii and A. adscendens (crosses). B. A. latifolius (solid triangles) and possible hybrids between A. latifolius and A. tessmannii (crosses). C. A. microcarpus (solid triangles) and A. cf. microcarpus (open triangles). D. The location in South America of the map section shown inA–C. Elevation is depicted in A–C as follows: 0–1000 m as white; 1001–2000 m, 2001–3000 m, 3001–4000 m, and>4000 m as progressively darker shades of grey.

FIGURE 6. Adelobotrys microcarpus Schulman, sp. nov. A habit showing abaxial (lower- and uppermost leaves) and adaxial sides of laminae, and inflorescence; B smaller (episepalous) stamen; C larger (epipetalous) stamen and petal; D bud showing cylindrical hypanthium, shallowly 5-lobed calyx, and absence of calyx teeth; E fruiting hypanthium with persistent calyx and clear constriction at torus; F remains of old, disintegrated hypanthium (fruit removed) exhibiting costae that are not apically connected by a circular vascular strand, also shown are the strongly elongated branches of the partial inflorescence with scars of fallen-off fruit; G seed with point of attachment to placenta marked; H close-up view of serrulate-setulose lamina margin from abaxial side; J malpighian hairs from lamina abaxial side (lower three from lamina proper, upper three from vein) with point of attachment marked on two hairs. Drawn by R. Ilmanen from J. J. Wurdack 2070. Scalebars: for A 5 cm, B–C and D–F 5 mm, G and J 0.5 mm, H 2 mm.

FIGURE 7. The occurrence of three species of Adelobotrys in soils of different soil cation concentration in four regions of Amazonian lowland rainforests (Juruá in central Brazil, Yasuní in Ecuador, Loreto in northern Peru, and Madre de Dios in southern Peru). Each open black circle indicates the natural logarithm of the sum of soil cation concentration (Ca+K+Mg+Na cmol(+)/kg) in one inventory transect (see Fig. 1 for locations of transects). The coloured circles indicate the soil cation concentration of an inventory transect where the species have been observed to occur. The box plot summarises, over all regions, the soil variation observed in those inventory transects where the species was encountered. The whiskers of the box embrace the minimum and maximum, the width of the box tells the limits of the first and third quartile, and the vertical line within the box shows the median of the natural logarithm of the sum of cation concentration. Orange colour stands for A. latifolius, green for A. microcarpus, and violet for A. tessmannii.