diff --git a/data/03/E6/A4/03E6A435FFA0FFE3C0BD74F1ABDB59C5.xml b/data/03/E6/A4/03E6A435FFA0FFE3C0BD74F1ABDB59C5.xml deleted file mode 100644 index f9aaf4456ac..00000000000 --- a/data/03/E6/A4/03E6A435FFA0FFE3C0BD74F1ABDB59C5.xml +++ /dev/null @@ -1,249 +0,0 @@ - - - -Leonardoxa africana (Leguminosae: Caesalpinioideae): a complex of mostly allopatric subspecies - - - -Author - -Mckey, Doyle B. -Centre d’Écologie Fonctionnelle et Évolutive (UPR CNRS 9056), Université Montpellier II, 1919 route de Mende, 34293 Montpellier cedex 05, France. -mckey@cefe.cnrs-mop.fr - -text - - -Adansonia - - -2000 - -3 - - -2000-06-03 - - -22 - - -1 - - -71 -109 - - - -journal article -http://doi.org/10.5281/zenodo.4605796 -1639-4798 -4605796 -933B60E6-57E6-4EB4-ACA8-D871ABD9F36A - - - - - - -Leonardoxa africana -subsp. -gracilicaulis - - - - -McKey, -subsp. nov. -(= group 1) - - - - -Haec subspecies -Leonardoxae africanae - -subsp. -rumpiensi - -arcte atque -L. africanae - -subsp. -letouzeyi - -et - -subsp. -africanae - - -nonnihil remotior affinis, sed ab ea glandulis nectariferis crateriformibus ad foliola proximalia nullis ad duabus atque foliorum expansione simultanea ramunculi ad nodos terminales duos ad quattuor, ab omnibus ramunculorum juvenilium internodiis non inflatis atque racemis ad ramunculos diametro -5-10 mm -axillaribus distinguitur. - - - - -TYPUS -. — - -Letouzey -10201, - -Cameroon -, -Southern Province -, -Colline Ongongondje -near -Akonekye -, - -15 km -NW Ambam - -, ca. - -700 m - -, - -23 Mar. 1970 - -(holo-, - -P; iso-, BR, K, WAG, YA). - - -Tree to -25 m -tall. Trunk up to -50-60 cm -diam. Young shoots produced in flushes of (1-)2-4 internodes and leaves at each branch tip; internodes of young twigs (1.1-)3.7(-7) cm long, not modified as myrmecodomatia, not swollen; twigs (2-)2.3(-3.1) mm diam. at apex tapering to ca. -1- 2 mm -diam. at base of internode. Prostoma lacking. Leaves 2-3-jugate; leaflets elliptic, falcate, apex acuminate to long-acuminate, the smallest ones (proximal pair) (9.6-)10.9(-15) cm long and (3-)4(-5) cm broad, largest ones (the distal ones in 2-jugate leaves, median ones in 3-jugate leaves) (12-)15(-26) cm long and (4-)5.5(-9.5) cm broad. Nectary glands small (smooth gland surface oblong, about 0.5 × -0.75 mm -), mostly restricted to proximal leaflets, where they are variable in number (0-2). Racemes very short (axis to -3 cm -long), with 25-40 flowers, axillary on small twigs of 2-3 (rarely 5-10) mm diam. ( -Fig. 8 -). Flowers -1.6-1.8 cm -in diam; pedicels -2-4 mm -long; calyx white to pale lilac, sepals -0.5-0.8 cm -long; petals lilac to violet, -0.6-1 cm -long. Ovules 4-5. Fruit oblong-obovate, ca. -10 cm -long, -4 cm -broad. - - - -Leonardoxa africana -subsp. -gracilicaulis - -is distinguished from the three other subspecies of the - -L. africana - -complex by the absence of swollen internodes and prostoma (the subspecies - -gracilicaulis - -is not a myrmecophyte: -Fig. 6A -). Foliar nectary glands are present, but are smaller than those of the myrmecophytic subspecies ( -Fig. 5A -). This subspecies also differs from the others in floral characters. Flowers are white and lilac in colour, rather than pink or red, the most frequent colours in the myrmecophytes. Also, inflorescences are usually axillary on small twigs in this subspecies ( -Fig. 8A -). Cauliflory is very frequent in the myrmecophytes, but in field populations and herbarium specimens studied by us, cauliflory has been recorded in the non-myrmecophytic subspecies only at the summit of Nta Ali ( -Table 6 -), in a population that may be affected by hybridisation with - -L. africana -subsp. -letouzeyi - -(see below). Trees of this subspecies, especially in hilltop (piedmont and lower submontane) habitats where it is most common, reach larger size than is ever attained by any of the myrmecophytes. - - - - -DISTRIBUTION. — This subspecies includes all the specimens collected from -Gabon -, -Equatorial Guinea -, and southern -Cameroon -from the vicinity of Yaoundé southward and east of the coastal plain ( -Fig. 4 -). In southern -Cameroon -, this subspecies is characteristically abundant on summits of hills, such as Nkolebengue ( -THOMAS & THOMAS 1993 -) and Zingui (this study). Near Yaoundé, - -L. africana -subsp. -gracilicaulis - -is one of the most frequent trees in submontane forest from -1,000 m -elevation upwards ( -ACHOUNDONG 1985 -, -1996 -). The nonmyrmecophytic - -L. africana - -abundant in forest on the summit of Nta Ali ( -ACHOUNDONG 1995 -) are also tentatively ascribed to this subspecies.This subspecies is much less frequent in lowland forests, where it is highly localised along streams and patchily distributed. (Within its range, lowland habitats are less constantly humid than hilltop and submontane habitats.) In the coastal plain, where the highly specialised myrmecophyte - -L. africana -subsp. -africana - -predominates (see below), - -L. africana -subsp. -gracilicaulis - -has been collected from two inselbergs (Colline Ngwon, -38 km -E of Kribi: -Letouzey 9371 -; Nkoltsia, -35 km -from the coast, -Villiers 807 -). Near the latter site, it also occurs in lowland forest, near the village Nkoloboundé along the stream Bounde, where it occurs together, interspersed in the same forests, with the specialised myrmecophyte - -L. africana -subsp. -africana - -. This is so far the only site where the two are known to cooccur.In forest along two other streams nearby,only - -L. africana -subsp. -africana - -was present. The two subspecies may co-occur elsewhere, though. The collection -Mpom 303 -, from -14 km -SE of Douala on the old Douala-Edea road, includes sheets of - - - - \ No newline at end of file diff --git a/data/03/E6/A4/03E6A435FFA0FFE3C0BD74F1ADDC5AB4.xml b/data/03/E6/A4/03E6A435FFA0FFE3C0BD74F1ADDC5AB4.xml new file mode 100644 index 00000000000..62beb449013 --- /dev/null +++ b/data/03/E6/A4/03E6A435FFA0FFE3C0BD74F1ADDC5AB4.xml @@ -0,0 +1,833 @@ + + + +Leonardoxa africana (Leguminosae: Caesalpinioideae): a complex of mostly allopatric subspecies + + + +Author + +Mckey, Doyle B. +Centre d’Écologie Fonctionnelle et Évolutive (UPR CNRS 9056), Université Montpellier II, 1919 route de Mende, 34293 Montpellier cedex 05, France. +mckey@cefe.cnrs-mop.fr + +text + + +Adansonia + + +2000 + +3 + + +2000-06-03 + + +22 + + +1 + + +71 +109 + + + +journal article +10.5281/zenodo.4605796 +1639-4798 +4605796 + + + + + + +Leonardoxa africana +subsp. +gracilicaulis +McKey + +, + +subsp. nov. + +(= group 1) + + + + + +Haec subspecies +Leonardoxae africanae + +subsp. +rumpiensi + +arcte atque +L. africanae + +subsp. +letouzeyi + +et + +subsp. +africanae + + +nonnihil remotior affinis, sed ab ea glandulis nectariferis crateriformibus ad foliola proximalia nullis ad duabus atque foliorum expansione simultanea ramunculi ad nodos terminales duos ad quattuor, ab omnibus ramunculorum juvenilium internodiis non inflatis atque racemis ad ramunculos diametro +5-10 mm +axillaribus distinguitur. + + + + + + +TYPUS. — + +Letouzey +10201 +, + +Cameroon +, +Southern Province +, +Colline Ongongondje +near +Akonekye +, + +15 km +NW + +Ambam +, ca. + +700 m + +, + +23 Mar. 1970 + +( +holo- +, +P +; + + +iso- +, +BR +, +K +, +WAG +, +YA +). + + + + + +Tree to +25 m +tall. Trunk up to +50-60 cm +diam. Young shoots produced in flushes of (1-)2-4 internodes and leaves at each branch tip; internodes of young twigs (1.1-)3.7(-7) cm long, not modified as myrmecodomatia, not swollen; twigs (2-)2.3(-3.1) mm diam. at apex tapering to ca. +1- 2 mm +diam. at base of internode. Prostoma lacking. Leaves 2-3-jugate; leaflets elliptic, falcate, apex acuminate to long-acuminate, the smallest ones (proximal pair) (9.6-)10.9(-15) cm long and (3-)4(-5) cm broad, largest ones (the distal ones in 2-jugate leaves, median ones in 3-jugate leaves) (12-)15(-26) cm long and (4-)5.5(-9.5) cm broad. Nectary glands small (smooth gland surface oblong, about 0.5 × +0.75 mm +), mostly restricted to proximal leaflets, where they are variable in number (0-2). Racemes very short (axis to +3 cm +long), with 25-40 flowers, axillary on small twigs of 2-3 (rarely 5-10) mm diam. ( +Fig. 8 +). Flowers +1.6-1.8 cm +in diam; pedicels +2-4 mm +long; calyx white to pale lilac, sepals +0.5-0.8 cm +long; petals lilac to violet, +0.6-1 cm +long. Ovules 4-5. Fruit oblong-obovate, ca. +10 cm +long, +4 cm +broad. + + + +Leonardoxa africana +subsp. +gracilicaulis + +is distinguished from the three other subspecies of the + +L. africana + +complex by the absence of swollen internodes and prostoma (the subspecies + +gracilicaulis + +is not a myrmecophyte: +Fig. 6A +). Foliar nectary glands are present, but are smaller than those of the myrmecophytic subspecies ( +Fig. 5A +). This subspecies also differs from the others in floral characters. Flowers are white and lilac in colour, rather than pink or red, the most frequent colours in the myrmecophytes. Also, inflorescences are usually axillary on small twigs in this subspecies ( +Fig. 8A +). Cauliflory is very frequent in the myrmecophytes, but in field populations and herbarium specimens studied by us, cauliflory has been recorded in the non-myrmecophytic subspecies only at the summit of Nta Ali ( +Table 6 +), in a population that may be affected by hybridisation with + +L. africana +subsp. +letouzeyi + +(see below). Trees of this subspecies, especially in hilltop (piedmont and lower submontane) habitats where it is most common, reach larger size than is ever attained by any of the myrmecophytes. + + + + +DISTRIBUTION. — This subspecies includes all the specimens collected from +Gabon +, +Equatorial Guinea +, and southern +Cameroon +from the vicinity of Yaoundé southward and east of the coastal plain ( +Fig. 4 +). In southern +Cameroon +, this subspecies is characteristically abundant on summits of hills, such as Nkolebengue ( +THOMAS & THOMAS 1993 +) and Zingui (this study). Near Yaoundé, + +L. africana +subsp. +gracilicaulis + +is one of the most frequent trees in submontane forest from +1,000 m +elevation upwards ( +ACHOUNDONG 1985 +, +1996 +). The nonmyrmecophytic + +L. africana + +abundant in forest on the summit of Nta Ali ( +ACHOUNDONG 1995 +) are also tentatively ascribed to this subspecies.This subspecies is much less frequent in lowland forests, where it is highly localised along streams and patchily distributed. (Within its range, lowland habitats are less constantly humid than hilltop and submontane habitats.) In the coastal plain, where the highly specialised myrmecophyte + +L. africana +subsp. +africana + +predominates (see below), + +L. africana +subsp. +gracilicaulis + +has been collected from two inselbergs (Colline Ngwon, +38 km +E of Kribi: +Letouzey 9371 +; Nkoltsia, +35 km +from the coast, +Villiers 807 +). Near the latter site, it also occurs in lowland forest, near the village Nkoloboundé along the stream Bounde, where it occurs together, interspersed in the same forests, with the specialised myrmecophyte + +L. africana +subsp. +africana + +. This is so far the only site where the two are known to cooccur.In forest along two other streams nearby,only + +L. africana +subsp. +africana + +was present. The two subspecies may co-occur elsewhere, though. The collection +Mpom 303 +, from +14 km +SE of Douala on the old Douala-Edea road, includes sheets of both + +L. africana +subsp. +gracilicaulis + +(in P and YA) and of + +subsp. +africana + +(in BR). + + + + +ECOLOGY. — This subspecies is not a myrmecophyte, but is “myrmecophilic,” attracting many different species of tree-dwelling ants to its foliar nectar. Some of these (e.g., + +Tetramorium aculeatum + +, + +Oecophylla longinoda + +) establish nests on leaf surfaces, but no ants nest in twigs. Ant-exclusion experiments found no effect of these opportunistically attracted ants in protecting young leaves of the plant against herbivores ( +GAUME 1998 +). This subspecies was termed “ + +L. gracilicaulis + +” by +MCKEY (1991) +and “ + +L. africana + +taxon T1” by +CHENUIL & MCKEY (1996) +and +BROUAT et al. (1998) +. + + + + + +PARATYPES +. — +CAMEROON +, +Central-South Province +: +Bitye +, + +Bates +1660 + +( +MO +) + +; + +Nkomnyat +, + +4 km +N + +Ekekam +, + +Dang +651 + +( +P +, +YA +) + +; + +hill near +Mt. Febe +, +Yaoundé +, + +Manning +2157 + +( +MO +) + +; + +Colline Ngwon +, + +38 km +E + +Kribi +, + +Letouzey +9371 + +( +P +, +WAG +, +YA +) + +; + +Evelessi +, + +30 km +W + +Sangmelima +, + +Letouzey +10158 + +( +BR +, +K +, +P +, +WAG +, +YA +) + +; + + +20 km +NE + +Yaoundé +, + +Mildbraed +8065 + +( +K +) + +; + +Makak +, + +SRFCam +/ +1141 + +( +P +, +YA +) + +; + +Lolodorf +, + +Staudt +427 + +( +G +, +K +) + +; + +colline +Nkoltsia +, + +Villiers +807 + +( +P +) + +; + + +5 km +S + +Mbalmayo +, + +W. de Wilde +1862 + +( +BR +, +K +, +MO +, +P +, +WAG +, +YA +) + +; + +Bipindi +, + +Zenker ed. Weigel +37 + +( +G +, +US +, +WAG +) + +, + + +Zenker +s.n + +. ( +BR +, +P +), + +1696 + +( +K +, +WAG +), + +1698 + +( +BR +), + +2255 + +( +K +), + +2255b + +( +BR +, +G +, +K +, +P +, +WAG +, +Z +), + +2303a + +( +K +, +P +), + +2989a + +( +BR +, +K +, +P +, +WAG +), + +3747 + +( +MO +), + +4545 + +( +BR +, +K +), + +4972 + +( +BR +, +K + +); + + +3km +WNW + +Bipindi +, + +Manning +1397 + +( +MO +) + +. + +Littoral Province +: + +14 km + +from +Douala +on the old road to +Edea +, + +Mpom + +303 + + +in part ( +P +, +YA +) + +. — + +GABON +, +Province Estuaire +: near +Libreville +, + +Klaine +3361 + +( +P +) + +; + +Sibange-Farm +, +Munda +, + +Soyaux +104 + +( +K +, +P +) + +; + +ca. + +15 km + +along road from +Assok +to hydroelectric power plant in the +Tchimbele River +, ca. + +600 m + +, + +Breteler +& +de Wilde +226 + +( +WAG +) + +. + +Province Woleu-Ntem +: region of +Oyem +, between the +Ogouée R +. and +Cameroon +, + +Le Testu +s.n. + +( +BR +, +K +, +MO +) + +, +9079 +(BR, P); + +Assoc Ngoum +, + +Le Testu +8995 + +( +P +, +BR +) + +. + +Province Ogouée-Ivindo +: +Mambanza +, between +Gazi +and +Dembona +, + +Le Testu +8888 + +( +B +, +K +, +MO +) + +; + +ca. + +24 km + +along +SOMIFER +road towards +Makokou +, elev. + +950-1000 m + +, + +Breteler +& +de Wilde +704 + +( +WAG +) + +; + +Village Manyal +, +Bord Ivindo +, + +Eckendorff +128 + +( +P +) + +. — + +EQUATORIAL GUINEA +: +Nkolentangun +, +Nscha’bat +(undetermined locality), + +Tessmann +284a + +( +K +) + +. — + +CONGO REPUBLIC +( +BRAZZAVILLE +): +Mbouamo +( +Sangha +) forêts (undetermined locality), + +Chevalier +s.n. + +( +P +) + +. + + + + \ No newline at end of file diff --git a/data/03/E6/A4/03E6A435FFA1FFDCC0BD763DA8B259C4.xml b/data/03/E6/A4/03E6A435FFA1FFDCC0BD763DA8B259C4.xml new file mode 100644 index 00000000000..5b9a71532e3 --- /dev/null +++ b/data/03/E6/A4/03E6A435FFA1FFDCC0BD763DA8B259C4.xml @@ -0,0 +1,325 @@ + + + +Leonardoxa africana (Leguminosae: Caesalpinioideae): a complex of mostly allopatric subspecies + + + +Author + +Mckey, Doyle B. +Centre d’Écologie Fonctionnelle et Évolutive (UPR CNRS 9056), Université Montpellier II, 1919 route de Mende, 34293 Montpellier cedex 05, France. +mckey@cefe.cnrs-mop.fr + +text + + +Adansonia + + +2000 + +3 + + +2000-06-03 + + +22 + + +1 + + +71 +109 + + + +journal article +10.5281/zenodo.4605796 +1639-4798 +4605796 + + + + + +3. + +Leonardoxa africana +subsp. +rumpiensis +McKey + +, + +subsp. nov. + +(= group 2) + + + + + +Haec subspecies +Leonardoxae africanae +subsp. letouzeyi +perarcte affinis, sed ab ea glandulis nectariferis crateriformibus majoribus ac quoad numerum plus constantibus (ad foliola proximalia plerumque nullis ad duabus) atque foliolis utroque rhachidis latere plerumque duobus distinguitur. + + + + + +TYPUS. — + +Letouzey +14577 + +, +Cameroon +, +Southwest Province +, +Dikome Balue +, +4°55’N +, +9°15’E +, + +35 km +NNW + +Kumba +, + +1200 m + +, + +24 Mar. 1976 + +( +holo- +, +P +; + + +iso- +, +K +, +YA +). + + + + + +Small tree, to +14 m +tall. Trunk to +35 cm +diam. Young shoots produced in flushes consisting of (1-)2-3 internodes plus young leaves at each branch tip (in adult trees) ( +Fig. 7 +). Internodes of young twigs (3-)5.5(-11) cm long, modified as myrmecodomatia, swollen, with thick pith; twigs (3-)4(-5) mm diam. at apex tapering to ca. +2-3 mm +diam. at base of internode; enlarged pith extending throughout section of twig produced in a single growth flush; ant-cavities each 2-3 internodes long (in adult trees). Swollen internodes absent in seedlings, present only in plants ≥ +50 cm +tall. Prostoma of ant-domatia at apex of terminal internode of each flush of growth, opposite leaf insertion, roughly spherical in shape. Leaves 2 (-3)-jugate ( +Fig. 9 +). Leaflets elliptic, falcate, apex acuminate to long-acuminate, smallest ones (proximal pair) (6.5-)16(-28) cm long and (2.7-) 6.5(-11) cm broad, largest leaflets (second pair) (7-)17.5(-30) cm long and (3-)7(-12) cm broad. Nectary glands of intermediate size (smooth gland surface oblong, about 1 × +0.75 mm +, not surrounded by a raised ring of tissue), usually 2(0-5) on proximal leaflets ( +Fig. 5 +), (0-)1(-2) on other leaflets. Racemes short (axis +3-5 cm +long), with> 40-60 flowers, axillary on twigs +4-5 mm +diam., also sometimes cauliflorous (on trunk up to +10 cm +diam.). Flowers +1.8-3 cm +diam.; pedicels +0.3-0.5 cm +long; calyx colour unknown, sepals +0.8-1.1 cm +long; petals mauve-pink to purple, +0.9-1.3 cm +long. Ovule number unknown. Size of mature fruit unknown. + + +This subspecies is also a myrmecophyte; all trees possess swollen hollow internodes occupied by ants. As mentioned above, + +L. africana +subsp. +rumpiensis + +shares a number of traits with + +subsp. +letouzeyi + +that distinguish these two from + +subsp. +africana + +. It is distinguished from + +subsp. +letouzeyi + +by its usually 2-jugate leaves ( +Fig. 9 +), the smaller size of its leaflets and stems, the constancy in nectary number, and its restriction to piedmont and submontane forest (above +500 m +and probably more commonly above +800 m +). + + + + +DISTRIBUTION. — This taxon occurs in the Rumpi Hills ( +Figs. 3 +, +4 +), where it was studied in 5 field sites. Herbarium specimens from three other sites in the Rumpi Hills (see below) also appear to belong here. Material of myrmecophytic + +L. africana + +collected from submontane forest on another hill in the Southwest Province ( +Letouzey 13632 +[K, P, YA], from a +960 m +hill +55 km +southwest of Mamfe) is tentatively placed in this subspecies. + + + + +ECOLOGY. — This myrmecophyte, in contrast to the two preceding subspecies, is restricted to submontane forest (and lowland forest transitional to it: “piedmont” [ +THOMAS 1995 +]). As in the other myrmecophytes, internodes are hollowed and occupied by ants, and each adult tree is occupied by a single ant colony. In contrast to the two preceding subspecies, each of which is principally associated with one (or two) ant species, a diverse array of ant species occupy trees in populations of the subspecies +rumpiensis +(D. MCKEY and R. SNELLING, unpubl. results). Nature of the interactions with ants has not been studied in this subspecies. This subspecies was termed “ + +L. africana + +taxon T2” by +CHENUIL & MCKEY (1996) +. + + + + + +PARATYPES +. — +CAMEROON +, +Southwest Province +: +Steep +hillside, + +400-850 m + +, + +5 km +W + +of +Esukutang Village +, + +D.W +. + +Thomas et al. +7988 + + +( +MO +); + + +forest, + +500 m + +, near + +Masaka-Batanga + +, + +D. +W +. + +Thomas & +Namata +7746 + + +( +MO +); + + +forest, + +500 m + +, near +Meta village +, + +Nemba +et al. +753 + +( +MO +); + + +colline, + +960 m + +(près côte 897) +E +of piste +Abakpa-Mbiofong +, + +55km +SW + +Mamfe +, + +Letouzey +13632 + +( +K +, +P +, +YA +) + +. + + + + \ No newline at end of file diff --git a/data/03/E6/A4/03E6A435FFA2FFDDC26977D7AAF15E8E.xml b/data/03/E6/A4/03E6A435FFA2FFDDC26977D7AAF15E8E.xml new file mode 100644 index 00000000000..e28c8ad3cb4 --- /dev/null +++ b/data/03/E6/A4/03E6A435FFA2FFDDC26977D7AAF15E8E.xml @@ -0,0 +1,412 @@ + + + +Leonardoxa africana (Leguminosae: Caesalpinioideae): a complex of mostly allopatric subspecies + + + +Author + +Mckey, Doyle B. +Centre d’Écologie Fonctionnelle et Évolutive (UPR CNRS 9056), Université Montpellier II, 1919 route de Mende, 34293 Montpellier cedex 05, France. +mckey@cefe.cnrs-mop.fr + +text + + +Adansonia + + +2000 + +3 + + +2000-06-03 + + +22 + + +1 + + +71 +109 + + + +journal article +10.5281/zenodo.4605796 +1639-4798 +4605796 + + + + + +2. + +Leonardoxa africana +subsp. +letouzeyi +McKey + +, + +subsp. nov. + +(= group 3) + + + + + +Haec subspecies +Leonardoxae africanae + +subsp +. africanae + + +arcte affinis, sed ab ea medullae loculo per internodia duo triave continuo, foliorum expansione simultanea ramunculi ad nodos terminales duos tresve, glandulis nectariferis crateriformibus minoribus ac quoad numerum plus variabilibus (ad foliola proximalia plerumque nullis ad duabus) atque foliolis usque ad +30 cm +longis utroque rhachidis latere saepe quattuor distinguitur. + + + + + + +TYPUS. — + +D.W. Thomas +et al. +7480 + +, +Cameroon +, +Southwest Province +, roadside forest +S +of +Baro Village +, +5º14’N +, +9º15’E +, + +250 m + +, + +31 Mar. 1988 + +( +holo- +, +MO +; + + +iso- +, +WAG +). + + + + + +Small tree, (rarely) to +14 m +tall. Trunk to +30 cm +diam. Young shoots produced in flushes consisting of (1-)2-3 internodes plus young leaves at each branch tip (in adult trees). Internodes of young twigs (2.8-)7(-13) cm long, modified as myrmecodomatia, swollen, with thick pith; twigs (4-)5.5(-6.1) mm diam. at apex tapering to ca. +2.5-4 mm +diam. at base of internode; enlarged pith extending throughout section of twig produced in a single growth flush; ant-cavities each 2-3 internodes long (in adult trees). Swollen internodes absent in seedlings, present only in plants ≥ +60 cm +tall. Prostoma of ant-domatia at apex of terminal internode of each flush of growth, opposite leaf insertion, roughly spherical in shape. Leaves (2-3-)4(-5)-jugate. Leaflets elliptic, falcate, apex long-acuminate, proximal ones (13.2-)17(-22) cm long and (4.5-)6(-8) cm broad, largest leaflets (next-to-distal pair) (14.5-) 21(-32) cm long and (5-)7.5(-12) cm broad. Nectary glands absent from some individuals; when present, appearing intermediate in size (smooth gland surface oblong, about 1 × +0.75 mm +, not surrounded by a raised ring of tissue) and few in number, usually 0-2 (very rarely up to 5) on proximal leaflets and usually absent on other leaflets. Racemes short (axis +3-5 cm +long), with 40-60 flowers, axillary on twigs of up to +1 cm +diam., also sometimes cauliflorous. Flowers +1.8-3 cm +diam.; pedicels +0.3-0.5 cm +long; calyx pale lilac, sepals +0.8-1.1 cm +long; petals usually violet, sometimes pink, +0.9-1.3 cm +long. Ovules 4-5. Fruit to +15 cm +long, +5 cm +broad. + + +The most distinctive trait of + +L. africana +subsp. +letouzeyi + +is the large size of its stems and leaves ( +Fig. 9C +). Young twigs are much more robust than those of corresponding age of plants of any of the other subspecies, leaflets are larger, and 4- jugate leaves predominate on most adult trees, in contrast to the 2-or 3-jugate leaves which characterise the other subspecies. Another trait distinguishing the subspecies + +letouzeyi + +from all other subspecies is the great variation among individual trees of the same population in the number of foliar nectaries. This subspecies shares with + +subsp. +rumpiensis + +several traits that together distinguish them from + +subsp. +africana + +: (1) Antcavities are first produced in juveniles over +50 cm +tall; (2) ant-cavities in adult trees extend over 2-3 internodes, corresponding to the units of growth produced in flushes; (3) vegetative growth is synchronised over the crown of the tree; (4) the prostoma is roughly spherical in shape ( +Fig. 6C +). + + + + +Fig. 9. — Typical leaves of the + +Leonardoxa africana + +complex, all drawn at same scale. Bar = 10 cm. +A +, + +L. africana +subsp. +gracilicaulis + +; +B +, + +L. africana +subsp. +rumpiensis + +; +C +, + +L. africana +subsp. +letouzeyi + +; +D +, + +L. africana +subsp. +africana + +. + + + + +DISTRIBUTION. — This subspecies occurs in the wettest forests of Africa, the lowland forests near the Bight of Biafra ( +Figs. 3 +, +4 +). It is found from Uwet Division, Calabar Province, +Nigeria +, through Korup National Park on Cameroon’s border with +Nigeria +, and throughout the Mamfe Plain and +Cross River +area. Its range extends northward at least to the Takamanda Forest Reserve north of Mamfe. The eastern limit of its distribution is uncertain. + + + + +ECOLOGY. — This subspecies is a myrmecophyte in which swollen stems may first appear in juveniles about +60 cm +tall (but more commonly in saplings +1 m +or taller). While saplings are associated with small colonies of any of several species of twig-nesting ants, mature individuals are occupied solely by the formicine ant + +Aphomomyrmex afer + +, which protects young leaves of the host tree against insect herbivores ( +GAUME 1998 +; +GAUME & MCKEY 1998 +). This subspecies was termed “ + +L. letouzeyi + +” by +MCKEY (1991) +and “ + +L. africana + +taxon T3” by C HENUIL & M C K EY (1996), +GAUME et al. (1998) +and +GAUME & MCKEY (1998) +. The ant occupants of this subspecies tend scale insects and mealybugs inside the hollow stems of the host ( +GAUME et al. 1998 +), in contrast to the ants associated with the subspecies + +africana + +, which never tend homopterans. (For the subspecies +rumpiensis +, more information is required on this point.) As argued by +GAUME et al. (1998) +, this dependence on homopterans is related to the low and variable number of foliar nectaries in this subspecies. + + + + + +PARATYPES +. — +NIGERIA +, +Uwet Division +: +Calabar River +, + +Latilo +35 + +( +BR +, +K +, +P +) + +; + +Oban +, + +Talbot +1440 + +( +K +) + +. — + +CAMEROON +, +Southwest Province +: +Bamenda Road +, +Mile 43 +from +Mamfe +, 1,500 ft ( + +460 m + +), + +Coombe +191 + +( +BR +, +K +) + +; + +Akpasang River +, +Korup National Park +, + +McKey +72 + +( +P +, +YA +) + +; + +Korup National Park +, between +Mana River +bridge and +Camp 1 +, + +Manning +1714 + +( +MO +) + +; + +disturbed forest near +Ndian Oil Palm Plantation +, + +50 m + +, + +D.W. Thomas +4269 + +( +BR +, +US +) + +; + +Takamanda Forest Reserve +, near +Matene +, + +170 m + +, + +D.W. Thomas +4527 + +( +MO +) + +. + + + + \ No newline at end of file diff --git a/data/03/E6/A4/03E6A435FFA6FFDEC269718BA8F05CC6.xml b/data/03/E6/A4/03E6A435FFA6FFDEC269718BA8F05CC6.xml new file mode 100644 index 00000000000..c76392aa13e --- /dev/null +++ b/data/03/E6/A4/03E6A435FFA6FFDEC269718BA8F05CC6.xml @@ -0,0 +1,868 @@ + + + +Leonardoxa africana (Leguminosae: Caesalpinioideae): a complex of mostly allopatric subspecies + + + +Author + +Mckey, Doyle B. +Centre d’Écologie Fonctionnelle et Évolutive (UPR CNRS 9056), Université Montpellier II, 1919 route de Mende, 34293 Montpellier cedex 05, France. +mckey@cefe.cnrs-mop.fr + +text + + +Adansonia + + +2000 + +3 + + +2000-06-03 + + +22 + + +1 + + +71 +109 + + + +journal article +10.5281/zenodo.4605796 +1639-4798 +4605796 + + + + + +1. + +Leonardoxa africana +(Baill.) Aubrév. subsp. +africana + +(= group 4) + + + + + +Tree to +14 m +tall. Trunk (rarely) to +40 cm +diam., usually under +15 cm +diam. Young shoots produced in flushes consisting of only a single internode and young leaf at each branch tip. Internodes of young twigs (2.5-)3.5-7.5(-9) cm long, modified as myrmecodomatia, swollen, with thick pith except at base; twigs +4.3-5.7 mm +diam. at apex tapering to ca. +2 mm +diam. at base of internode; ant-cavities each a single internode long. Swollen internodes present from the seedling stage ( +20 cm +tall) onward. Prostoma of ant-domatia at apex of internode opposite leaf insertion, elliptic. Leaves (2-)3(-rarely 4)-jugate. Leaflets elliptic, falcate, apex long-acuminate, the smallest leaflets (proximal pair) (7.8-)13(-15) cm long and (3.5-)5(-6) cm broad, the largest ones (distal pair in 2-jugate leaves, next-to-distal pair in 3- and 4-jugate leaves) (9.5-)15(-20) cm long and (4-)5.5(-7) cm broad; nectary glands large (smooth gland surface oblong, about 1 × +0.75 mm +, surrounded by a raised ring of tissue about +1mm +broad), consistently present on abaxial surface of each leaflet, with (2-)3-5 glands on each proximal leaflet, (1-)2(-3) glands on each median leaflet, and (0-)1-2 glands on each distal leaflet. Racemes to +4 cm +long, with 40-50 flowers, usually cauliflorous, borne on main trunk and on large branches, or ramiflorous on twigs of ca. +5 mm +diam. and larger. Flowers +1.8-2.2 cm +in diam.; pedicels +2-4 mm +long; calyx green or red, sepals +0.5-0.8 cm +long; petals violet, purple, mauve, pink or red, 0.6-1.0 cm long. Ovules 4-5. Fruit oblongobovate, +8-11 cm +long, +2.5-4 cm +broad. + + + +Fig. 4. — Geographic distribution of herbarium collections of the + +Leonardoxa africana + +complex examined in this study: + +L. africana +subsp. +gracilicaulis + += Group 1; + +L. africana +subsp. +rumpiensis + += Group 2; + +L. africana +subsp. +letouzeyi + += Group 3; + +L. africana +subsp. +africana + += Group 4. + + + + +Leonardoxa africana +subsp. +africana + +is a highly specialised myrmecophyte, differing from the other myrmecophytic subspecies in a number of traits: (1) Foliar nectaries ( +Fig. 5C +) are larger, more numerous, and less variable in presence/absence and number than in the other subspecies; (2) only a single leaf-bearing internode is produced in each flush of growth at a branch tip, so that each ant-cavity is a single internode long ( +Figs. 6B +, +7B +); (3) growth of different branch tips is not synchronised over the crown of the tree; (4) specialised swollen internodes appear much earlier (seedlings +20 cm +tall) in plant ontogeny than in the other subspecies; (5) the prostoma is elliptic-oblong rather than spherical in shape ( +Fig. 6D +). Also, while cauliflory on old branches, or even on the trunk, occurs in all three myrmecophytic subspecies, it appears to be especially frequent in + +subsp. +africana + +( +Fig. 8B +). + + + + +DISTRIBUTION. — + +Leonardoxa africana +subsp. +africana + +is restricted to coastal forests of +Cameroon +( +Figs. 3 +, +4 +). All the field sites are below + +80 m + +elevation. Judging from collection localities and notes on labels, this subspecies appears rarely to occur above about + + +100 m + +. + +The +northernmost known localities are from the +Southern Bakundu Forest Reserve +, and the southern boundary of its range appears to be near +Campo +, at the border with +Equatorial Guinea +. +Within +this coastal strip, it extends inland as far as +Bipindi +, on the eastern edge of the coastal plain. + + + + +Fig. 5. — Foliar nectaries of the + +Leonardoxa africana + +complex, all shown at the same scale: +A +, + +L. africana +subsp. +gracilicaulis + +(Zingui hill site); +B +, + +L. africana +subsp. +rumpiensis + +(Dikomé Balué site); +C +, + +L. africana +subsp. +africana + +(Akanga site). Nectaries of + +L. africana +subsp. +letouzeyi + +(not pictured) are similar in size to those of + +subsp. +rumpiensis + +. + + + + +Fig. 6. — Internodes of the + +Leonardoxa africana + +complex, showing variation in specialisations for housing ants. Bar = 2 cm in all photos. +A +, apex of internode (note point of attachment of the paripinnately compound leaf) of + +L. africana +subsp. +gracilicaulis + +(Mbalmayo site), showing absence of swelling in this non-myrmecophytic subspecies; +B, +longitudinal section of internode of + +L. africana +subsp. +africana + +(Bombé Bakundu site) showing the strongly swollen ant-domatium, which in this subspecies is a single internode long.— +C +and +D +, prostoma of two myrmecophytic subspecies, located in both cases at the apex of the internode opposite the point of leaf insertion; the diagonally cut structure at the top of each photo is the leaf rachis: +C, +circular prostoma of + +L. africana +subsp. +letouzeyi + +(Rengo Rock site); prostoma of + +L. africana +subsp. +rumpiensis + +(not shown) is roughly similar in size and shape; +D +, elongate prostoma of + +L. africana +subsp. +africana + +(Bombé Bakundu site). Both are on internodes produced by plants growing in a greenhouse at the University of Miami, on cuttings uninhabited by ants. The structures are not holes, but are scars produced by the spontaneous drying of the unlignified prostoma in mature internodes. + + + + +Fig. 7. — Examples of variation in the unit of growth within the + +Leonardoxa africana + +complex. +A +, a branch tip of + +L. africana +subsp. +rumpiensis + +(Dikome Balue site) showing two recently matured internodes and leaves produced in a flush of growth; these leaves are bright green (shiny in this photograph) and distinct from the older mature leaf (dark green, not shiny) produced in a previous growth flush; arrow marks the base of this flush. Bar = 10 cm. +B +, a branch tip of + +L. africana +subsp. +africana + +(Bombe Bakundu site) showing the one-leaf and internode unit of growth typical of all populations of this subspecies. Bar = 2 cm. + + + + +Fig. 8. — Variation in floral traits within the + +Leonardoxa africana + +complex. Bar = 2 cm in both photos. +A +, inflorescence axillary on a young twig (vertical object just behind the inflorescence) in + +L +. +africana +subsp. +gracilicaulis + +(Mbalmayo site); hypanthium white, corolla and filaments of stamens lilac, anthers yellow. +B +, trunciflorous inflorescences in + +L. africana +subsp. +africana +(Grand Zambi) + +; hypanthium and corolla pink. + + + + +ECOLOGY. — This subspecies, illustrated in +AUBRÉVILLE (1970) +, is the one that has attracted most attention as a myrmecophyte. The studies by +ELIAS (1980) +, +MCKEY (1984) +, +GAUME et al. (1997) +and +GAUME & MCKEY (1999) +all concern this subspecies. Internodes are swollen and hollow, and these ant-domatia are present already in seedlings +20 cm +tall (see +Fig. 2B +in +LÉONARD [1993] +). This subspecies is associated with two ant species that are completely restricted to this host, the formicine + +Petalomyrmex phylax + +and the myrmicine + +Cataulacus mckeyi +( +MCKEY 1984 +) + +. + +Petalomyrmex + +is a mutualist of the plant, protecting its young leaves against phytophagous insects ( +GAUME et al. 1997 +), while + +Cataulacus + +is a parasite, providing no protection to the plant and excluding + +Petalomyrmex + +from plants it occupies ( +GAUME & MCKEY 1999 +). This subspecies was termed “ + +L. africana + +taxon T4” by +CHENUIL & MCKEY (1996) +. + +Few observations of flower visitors have been made, but on a few occasions sunbirds were seen visiting the pink to red flowers, which in this subspecies at least seem to have a somewhat longtubular corolla, compared to flowers of the non-myrmecophytic subspecies (see below). + + + +ADDITIONAL SPECIMENS EXAMINED. — + +CAMEROON +, +Southwest Province +: +Southern Bakundu Forest Reserve +, + +E.W. Jones in Brenan +9499 + +( +K +, +P +), + +FHI +29511 + +(K), + +29672 + +(K), and + +41007 + +(K); + + +Southern Bakundu Forest Reserve +, + +1 km +W + +Bombe Bakundu +, + +Manning +1499 + +( +MO +); + + +Southern Bakundu Forest Reserve +, near +Kindonge Camp +, + +Manning +1517 + +( +MO +); + + +Kumba +(planted in forestry nursery), + +FHI +29369 + +( +K +). + + +Littoral Province +: + +24 km +E + +Douala +, + +Breteler +et al. +2580 + +( +BR +, +K +, +P +, +YA +); + + +Douala +, route +Razel +, + +Endengle +2105 + +( +P +, +YA +); + + +Douala +, forêt de +Yelfoume +, + +Fleury +, in herb. +Chevalier +33346 + +( +P +); + + + +14 km +E + +Douala +, +Douala-Edea Road +, + +Mpom + +303 + + +in part ( +BR +); + + +Lake Tissongo +, +Douala-Edea Forest Reserve +, + +McKey +& +Gartlan +120 + +( +K +), + + + +D.W. Thomas +167 + +( +YA +), + + + +Waterman +& +McKey +846 + +( +K +); + + +stream +Mangombe +near +Edea +, + +Mpom +343 + +( +YA +). + + +Central-South Province +: near +Bipaga II +, +km 40 +from +Kribi +to +Edea +, + +de +Kruif +994 + +( +WAG +); + + +Lolodorf Road +, + +12 km +E of + +Kribi +, + +Bos +3630 + +( +BR +, +WAG +, +YA +); + + + +5 km +N of + +Lolodorf +road, + +19 km +E of + +Kribi +, + +Bos +3913 + +( +BR +, +YA +); + + +Lolodorf Road +, + +18 km +E of + +Kribi +, + +Bos +4074 + +( +BR +, +YA +); + + +km 30 +from +Kribi +to +Ebolowa +, + +Bos +6233 + +( +WAG +); + + +km 45 +from +Kribi +to +Campo +, + +Bos +7315 + +( +BR +); + + + +60 km +S of + +Edea +, +S of +Mboke +, + +11 km +E of km 58 + +from +Edea +to +Kribi +, + +Leeuwenberg +5500 + +( +BR +, +K +, +MO +, +P +, +WAG +, +YA +); + + +colline +Nkoltsia +, + +Villiers +775 + +( +P +); + + +near +Bella +, + +Letouzey +4140 + +( +P +, +YA +); + + + +2 km + +from +Nkol-Bewa +, + +Mpom +227 + +( +P +, +YA +); + + +Edoudouma +, + +Mpom +283 + +( +YA +); + + +Bipindi +, + +Farron +7181 + +( +BAS +, +P +, +YA +), + +Zenker +, s.n. + +( +BR +, +P +), + +52 + +( +WAG +), + +1074 + +( +K +, +MO +, +P +, +WAG +), + +2967 + +( +BR +, +K +, +WAG +), + +4183 + +( +BR +, +K +, +P +), + +4495 + +( +BR +, +K +, +MO +). + + + + + \ No newline at end of file diff --git a/data/03/E6/A4/03E6A435FFA7FFDAC0BD75FEABDB5AF9.xml b/data/03/E6/A4/03E6A435FFA7FFDAC0BD75FEABDB5AF9.xml new file mode 100644 index 00000000000..ae2ebae5b93 --- /dev/null +++ b/data/03/E6/A4/03E6A435FFA7FFDAC0BD75FEABDB5AF9.xml @@ -0,0 +1,198 @@ + + + +Leonardoxa africana (Leguminosae: Caesalpinioideae): a complex of mostly allopatric subspecies + + + +Author + +Mckey, Doyle B. +Centre d’Écologie Fonctionnelle et Évolutive (UPR CNRS 9056), Université Montpellier II, 1919 route de Mende, 34293 Montpellier cedex 05, France. +mckey@cefe.cnrs-mop.fr + +text + + +Adansonia + + +2000 + +3 + + +2000-06-03 + + +22 + + +1 + + +71 +109 + + + +journal article +10.5281/zenodo.4605796 +1639-4798 +4605796 + + + + + + +Leonardoxa africana +(Baill.) Aubrév. + + + + + + +Adansonia, ser. 2, 8: 179 (1968). — + +Humboldtia africana +Baill., Hist. Pl. + +2: 99 (1870). + + + + +— + +Type: + +Mann +726 + +, +Cameroon +, +Littoral Province +, +Cameroon River +(= wouri estuary), 1861 ( +holo- +, +K +; + + +iso- +, +P +) + +. + + + + +The description of + +Leonardoxa + +by +LÉONARD (1993) +is still generally accurate, but must be amended in the following ways to take into account the variation described here: twigs with or without a fusiform, hollow infra-nodal swelling; inflorescences axillary on young twigs, ramiflorous, or cauliflorous. + + + + + +Key to the subspecies of + +Leonardoxa africana + + + +Some of the characters included in the key follow predictable ontogenetic patterns. In small saplings and in sucker shoots from cuttings, for example, leaflets are smaller, fewer leaflet pairs are present, and fewer nectaries occur on each leaflet than in mature trees. The descriptions here, unless otherwise stated, apply to mature trees. + + + +1. Internodes of young twigs not swollen or hollow, not inhabited by ants; leaflet apex short-acuminate; inflorescences axillary on small twigs up to about 5(-rarely 10) mm diameter; hypanthium white to pale lilac, corolla lilac ................................................................................................................ 4. + +subsp. +gracilicaulis + + + + + +1’. Internodes of young twigs (at least on all plants ≥ +1.5 m +tall) swollen, with thickened pith, hollowed out and inhabited by ants; leaflet apex long-acuminate; inflorescences usually ramiflorous (on twigs> +10 mm +diameter) or cauliflorous on large branches and trunk, rarely axillary on young twigs; hypanthium and corolla mauve to pink, purple or red ...................................................................................................................... 2 + + + + + + +2. Young growth produced in flushes of a single internode at each branch tip, pith chamber thus restricted to single-internode segments; specialised swollen internodes present at all stages of growth, from seedling ( +20 cm +tall) to adult; largest leaflets usually +13-15 cm +long, +4.5-5.5 cm +broad; (2-)3(-5) nectaries at base of each proximal leaflet ........................................................................................................................ 1. + +subsp. +africana + + + + + +2’. Young growth produced in flushes of 2-4 internodes at each branch tip, pith chamber thus continuous over 2-4 internodes; swollen internodes present only on plants ≥ +60 cm +tall, seedlings lacking myrmecodomatia; largest leaflets usually +16-23 cm +long, +6-8 cm +broad; nectaries at base of each proximal leaflet variable in number, sometimes absent.................................................................................................................................. 3 + + + + + + +3. Leaves often 4(3-5)-jugate; nectaries often absent at base of each proximal leaflet, when present usually 1(-2) in number; lowland rain forest up to +300 m +elevation ...................................................... 2. + +subsp. +letouzeyi + + + + + +3’. Leaves usually 2(2-3)-jugate; nectaries at base of each proximal leaflet present on almost all plants, usually +2 in +number; submontane rain forest in the Rumpi Hills (above +800 m +), and possibly other hills in southwestern +Cameroon +.................................................................................................................... 3. + +subsp. +rumpiensis + + +Figures 3 +and +4 +show the distribution of field sites and herbarium collections, respectively, of each of the four infraspecific taxa recognised here. + + + + + \ No newline at end of file diff --git a/data/03/E6/A4/03E6A435FFBEFFC1C0BD7753AB855CE7.xml b/data/03/E6/A4/03E6A435FFBEFFC1C0BD7753AB855CE7.xml new file mode 100644 index 00000000000..132c186471b --- /dev/null +++ b/data/03/E6/A4/03E6A435FFBEFFC1C0BD7753AB855CE7.xml @@ -0,0 +1,151 @@ + + + +Leonardoxa africana (Leguminosae: Caesalpinioideae): a complex of mostly allopatric subspecies + + + +Author + +Mckey, Doyle B. +Centre d’Écologie Fonctionnelle et Évolutive (UPR CNRS 9056), Université Montpellier II, 1919 route de Mende, 34293 Montpellier cedex 05, France. +mckey@cefe.cnrs-mop.fr + +text + + +Adansonia + + +2000 + +3 + + +2000-06-03 + + +22 + + +1 + + +71 +109 + + + +journal article +10.5281/zenodo.4605796 +1639-4798 +4605796 + + + + + + +THE + +LEONARDOXA AFRICANA + +COMPLEX + + + + + + +The range of + +Leonardoxa africana + +extends from +Gabon +and +Equatorial Guinea +northward through southwestern +Cameroon +and into extreme southeastern +Nigeria +. The species has long attracted attention as a myrmecophyte ( +BEQUAERT 1922 +; +MILDBRAED 1922 +; +SCHNELL & BEAUFORT 1966 +; +ELIAS 1980 +; +MCKEY 1984 +; +LÉONARD 1993 +). +SCHNELL & BEAUFORT (1966) +, however, noted that swollen-stem myrmecodomatia were present only in some of the specimens from +Cameroon +they examined and were consistently absent from specimens collected from elsewhere in the range of the species, notably +Gabon +. LETOUZEY, the botanist who collected most widely in +Cameroon +, also noted the geographic heterogeneity of + +L. africana + +in presence vs. absence of swollen internodes ( +LETOUZEY 1985 +; see also notes accompanying numerous specimens and unpublished field notes). + + +Field, greenhouse and herbarium studies reported on here demonstrate that + +L. africana + +is heterogeneous in this and many other characters. The variation concerns several traits that are obviously related to symbiosis with ants (ant-domatia, foliar nectaries), as well as other vegetative characters not directly related to ant-plant interactions (architectural differences related to rhythms of growth), and some floral characters (cauliflory, flower colour). + + +Some of the characters examined here merit special attention. Traits of myrmecodomatia have not often been used in taxonomic studies of groups including myrmecophytes, possibly because botanists have often mistakenly regarded these structures as galls, induced or altered in form by insects or other agents (see, for example, +SCHNELL & BEAUFORT 1966 +). + + +In + +Leonardoxa + +, there is variation among the taxa recognised here not only in presence or absence of ant-domatia, but also in several aspects of their structure, and in the timing of onset of expression of these specialised stem swellings in the ontogeny of the plant. + + +Traits related to the phenology of vegetative growth are likewise rarely considered in taxonomic studies. Rhythmic growth of shoots characterises all + +L. africana + +, but the taxa recognised here differ in the number of internodes produced in each spurt of growth (a difference with architectural consequences, evident on both living plants and herbarium specimens) and in the degree to which growth is synchronised among branches (evident only on living trees). Differences in all these traits persist when plants are grown in greenhouses without ants, in common gardens, and in the one known instance when two taxa occur together in the same habitat. Differences in these traits are genetically based and taxonomically informative. + + +Discontinuities in variation in these and other characters consistently reveal a pattern of four distinct taxa, mostly allopatric in distribution. These new taxa are here described as subspecies of + +L. africana + +, reflecting the opinion that these taxa are not yet completely differentiated and reproductively isolated species. Three of the four subspecies are myrmecophytes, with swollen stems that +form myrmecodomatia +and house symbiotic ants. The fourth is not a myrmecophyte but, like the other subspecies (as well as many other +Detarieae +), possesses foliar nectaries, which attract a diversity of non-resident ants. Experimental and observational field studies ( +MCKEY 1991 +; +GAUME et al. 1997 +, +1998 +; +GAUME & MCKEY 1998 +, +1999 +; +GAUME 1998 +) show that the four taxa recognised here are not only morphologically but also ecologically distinct. Although apparently only incompletely differentiated, they appear to be following independent evolutionary trajectories. This is most clearly illustrated by the coexistence in at least one site of the nonmyrmecophyte and the most highly specialised of the myrmecophytes (degree of specialisation being defined in terms of the number of new characters derived since divergence from the common ancestor). In sympatry, these two taxa remain morphologically and ecologically distinct, suggesting the existence of barriers to gene exchange. Molecular genetic studies (BROUAT et al., in press; C. BROUAT, unpublished data) confirm the distinctness of these coexisting populations and support the taxonomic separations made here on morphological grounds. These studies also suggest past or present gene flow between these taxa. + + + + \ No newline at end of file