AMARANTHUS L
69. BURSERACEAE
than pistil, the filaments villous near the middle. Fruits of 3-5 drupes to 17 mm long, with a medial ridge, green becoming red-orange or black at maturity; seeds ellipsoid, to ca 14 mm long. Croat 8441, 9508.
Occasional, in the forest; locally common, at least in the vicinity of Zetek Trail 250. Flowers in the late dry season (March and April). The fruits mature within 2 months.
The fruits are taken by many birds, including fly- catchers, motmots, thrushes, and chachalacas (Duke,
1968).
Guatemala to Panama and south to Brazil; Lesser Antilles. In Panama, known from tropical moist forest on BCI and from premontane wet forest in Panama (Cerro Campana). Reported from tropical wet and premontane rain forests in Costa Rica (Holdridge et al., 1971).
See Figs. 300 and 301.
69. BURSERACEAE
Trees, often with aromatic sap. Leaves alternate, pinnate, petiolate; leaflets entire; venation pinnate; stipules lack- ing. Flowers usually functionally unisexual (dioecious) but similar in appearance, actinomorphic, minute, in axillary or rarely terminal, cymose panicles or racemes;
calyx cupulate, 3-5-lobed; petals 3-5, free or rarely connate into a lobed tube; disk present; stamens usually 2 times the number of corolla lobes, in 2 whorls; anthers 2-celled, introrse, dehiscing longitudinally; ovary super- ior, 2-5-locular, 2-5-carpellate; placentation axile; ovules 2 per locule, anatropous; style 1; stigma 1, simple or in some cases 2-5-lobed. Fruits drupaceous, each a tardily dehiscent capsule or a drupe with 1-5 pyrenes (Trattin-
KEY TO THE SPECIES OF BURSERACEAE Petals united to form a tube at base:
Leaves conspicuously asperous, scabridulous on both surfaces; flowers 3-parted, reddish; fruits ovoid, ca 1 cm diam, indehiscent, the seeds (1)2 Trattinnickia aspera (Standl.) Swart Leaves not asperous, sparingly pubescent; flowers 4- or 5-parted, greenish-yellow; fruits broadly
turbinate to depressed-globose, more than 2.5 cm diam, the valves deciduous, the seeds usually 3 or more Tetragastris panamensis (Engler) O. Kuntze Petals free:
Leaves deciduous, the flowers appearing with new leaves; fruits ± 3-sided, the seed 1, white, 3-sided, indurate, remaining attached at base after valves fall; bark on trunk and larger branches papery, reddish-brown, often peeling Bursera simaruba (L.) Sarg.
Leaves not deciduous, the flowers concurrent with mature leaves; fruits or seeds not 3-sided, the seeds covered with a fleshy white aril, falling from fruit at maturity; bark on trunk and branches not papery, reddish-brown, not peeling:
Leaves and inflorescences glabrous; fruits glabrous; mature fruits usually obtuse to acute at both ends Protium panamense (Rose) I. M. Johnston Leaves and inflorescences pubescent (at least inflorescence branches; leaves glabrescent in P.
tenuifolium subsp. sessiliflorum); fruits pubescent (often scantily so at maturity); mature fruits obtuse to rounded at base:
Flowers 4-parted, pedicellate; petioles, rachises, and petiolules densely brownish-hirtellous, the leaf blades (at least below) sparsely hirtellous; fruits ± ovoid, mostly obtuse at base and at apex, the seed 1 Protium costaricense (Rose) Engler Flowers 5-parted, sessile or nearly so; petioles, rachises, and petiolules not densely
brownish-hirtellous, the leaf blades glabrous except for minute papillae; fruits usually rounded at both ends, 2-5-lobed, the seeds frequently 2-5 Protium tenuifolium Engler subsp. sessiliflorum (Rose) Porter
Fig. 300. Simarouba amara var. typica
Fig. 301. Simarouba amara var. typica
nickia); seeds 1 (rarely 2) per pyrene, lacking endosperm.
Members of the family may be confused with Anacar- diaceae (76) and Meliaceae (70), both of which have alternate, similarly compound leaves with leaflets fre- quently inequilateral at the base. The BCI species of Burseraceae can be distinguished from these other fami- lies by the presence of resin ducts in the bark, by the free stamens and single short style, and by the absence of pellucid dots. The sap of most species is resinous and has a characteristic aroma that is faintly turpentine-like but pleasant.
The pollination system is unknown, but the flowers are best suited to insect pollination.
The fruits are probably chiefly bird dispersed. Those of Protium and Tetragastris have white arillate seeds, which are displayed against the often bright-red inner carpel wall after one of the valves falls free. Seeds are often pendent, ideally suited to bird dispersal. The pyrenes or pits are very hard and probably pass through the digestive tract unharmed. The fruits are also taken by white-faced and spider monkeys (Oppenheimer, 1968), which eat the fleshy aril. Fruits of Tetragastris panamensis are taken by howler monkeys (Carpenter, 1934).
About 20 genera and 600 species; in the tropics.
BURSERA Jacq. exL.
Bursera simaruba (L.) Sarg., Card. & Forest 3:260. 1890
Naked Indian, Almacigo, Carate, Huechichi, Indio desnudo Dioecious or polygamodioecious tree, 5-25 m tall, to ca 40(100) cm dbh; bark coppery red, shiny, thin, peeling to expose green layer beneath; sap, at least of fruit, very aromatic. Leaves pinnate, deciduous, clustered at apex of branches, densely woolly when young to glabrate except on veins below in age; petioles to 14 cm long; leaflets 5-7 (9), ovate-elliptic to lanceolate-elliptic (terminal ones usually obovate), long-acuminate, inequilateral and ob- tuse to rounded at base, 4.5-14.5 cm long, 2.5-8 cm wide, entire. Flowers 3-5-parted, functionally unisexual, in axillary raceme-like panicles (sometimes appearing ter- minal before leaves appear), appearing ± with new leaves; calyx bowl-shaped, shallowly 5-lobed, the lobes acute to blunt; petals greenish-white, narrowly ovate, acute, cucullate, 2-3 mm long, spreading at anthesis, later recurved; stamens twice the number of and shorter than petals, those opposite the petals usually ± spreading, the alternate ones erect; pollen golden-yellow, covering all sides of anther at anthesis; pistillode ovoid, white, glabrous, scarcely longer than weakly lobed and undulate disk. Capsules drupaceous, ellipsoid, obtusely 3-sided, maturing reddish-brown, ca 1 cm long, a single valve fall- ing free at maturity to expose the pyrene, followed by both remaining valves falling off as a unit, leaving the pyrene attached at base; pyrene 1(2), 1-seeded, 3-angled, bony, lenticular-ovoid, white, ca 7 mm long. Croat 5325.
Occasional, but locally common, especially along the shore on the northwest side of the island and on Orchid Island. Flowers in the late dry and early rainy seasons (March to the middle of June), just before or during the onset of new leaves. Mature fruits may be seen through-
69.
BURSERACEAE/PROTIUM497
out much of the year, but most fruits mature during the late rainy or early dry seasons of the following year, generally after the plant has lost its leaves. Most trees are bare by February, beginning to put on new leaves and flowering by March or April. Allen (1956) reported that the trees are leafless throughout most of the dry season in Costa Rica.
Easily recognized by the reddish-brown, papery bark.
Southern Florida and northeastern Mexico to Colom- bia, Venezuela, and the Guianas; West Indies. In Panama, a characteristic species in tropical moist forest (Tosi, 1971), known all along the Pacific slope; known also from from premontane dry forest in Code, from tropical dry forest in Los Santos, Code, and Panama, and from pre- montane moist forest in the Canal Zone and Panama.
Reported from tropical wet forest in Costa Rica (Hol- dridgeetal, 1971).
PROTIUM Burm.f.
Protium costaricense (Rose) Engler, Nat. Pfl. ed. 2, 19a:414. 1931
P. salvozae Standl.
Functionally dioecious tree, to 11(25) m tall, the trunk usually less than 20 cm dbh; outer bark minutely rough- ened; inner bark reddish, forming minute, cloudy, viscid droplets; sap lacking typical burseraceous aroma; stems, petioles, rachises, inflorescence branches, and veins of leaflet pubescent, the trichomes stiff, erect, brownish, of varying lengths. Leaves pinnate; petioles 4-6.5 cm long, flattened on upper surface; leaflets (3) 5-7 (9), mostly oblong-elliptic, abruptly acuminate, obtuse to acute at base, slightly inequilateral, 6-17 cm long, 2.5-8 cm wide.
Flowers cream, 4-parted, in axillary panicles 1-13.5 cm long, often branched from near base; pedicels 1.5-3 mm long, puberulent; calyx bowl-shaped, broader than long, 1-2 mm long, shallowly lobed, the lobes acute to rounded;
petals ca 3 mm long, narrowly ovate, acute and weakly spreading at apex; stamens 8, arising from beneath a prominent disk, the disk weakly lobed, half as high as ovary in pistillate flowers (2 mm tall), as high as ovary in staminate flowers; filaments inflexed; anthers held directly over style; pollen white, tacky; ovary ovoid, appressed-pubescent, much shorter than stamens; style
± sessile; stigma 4-lobed. Capsules drupaceous, ± ovoid, 1.5-2 cm long, violet-purple to brownish, sparsely lenti- cellate, obtuse at both ends, the valves 2 (5), one falling free; pyrene 1, tan, 1-seeded, ca 1 cm long, falling free from funiculus (this ca 5 mm from apex) and suspended on a narrow band of tissue, enveloped in a white, sweet, fleshy aril displayed against the bright red inner valve surface. Croat 8262, 14926.
Rare, in the older forest along Zetek and Drayton trails.
Apparently flowers twice per season, once in February and early March, with the fruits maturing in March and April, and again in May and June, with the fruits devel- oping mostly in August and September.
Known from Costa Rica and Panama. In Panama, known only from tropical moist and premontane wet forests in the Canal Zone on the Atlantic slope; no doubt occurring on the Atlantic slope of western Panama also.
Fig. 304. Trattinnickia aspera
Protium panamense (Rose) I. M. Johnston, Contr.
Gray Herb. 70:72. 1924 Copa
Functionally unisexual tree, usually less than 12 m tall (to 40 m elsewhere), ± stilt-rooted at base, glabrous;
outer bark thin, smooth, the leaf scars on younger trees ca 5 cm broad; inner bark pinkish; sap with sweet strong odor typical of family. Leaves pinnate; petioles 6-13 cm long, flattened on upper surface, swollen at base; pet- iolules 1-4 (8) cm long, swollen at both ends; leaflets 3-7 (9), mostly ovate-lanceolate to oblong, bluntly short- acuminate, obtuse to rounded at base, 13-35 cm long, 6-9(16) cm wide, coriaceous. Inflorescences paniculate (rarely racemose), axillary or terminal, sometimes cauli- florous, mostly to 15 cm long; flowers greenish-white to greenish-yellow, 4 (5)-parted, with a strong, sweet aroma; pedicels to ca 4 mm long; calyx ± truncate or shallowly lobed, very short; petals 3 mm long, valvate,
± spreading at anthesis, papillose-puberulent on margins, the apex acute; stamens 8 (10), 2 mm long; filaments broadened below, recessed somewhat in fleshy yellowish disk; pistil conical, short; stigma simple, almost sessile, persistent in fruit. Capsules drupaceous, red at maturity, often in a large congested cluster, ellipsoid to ovoid, apiculate, shortly stipitate, to 2.8 cm long, the valves 2 or 4, unequal, the smaller falling free at maturity; pyrene 1(4), to ca 1.5 cm long, bearing 1 or 2 seeds, greenish, covered with a thick, white, fleshy, sweet mesocarp ca 3 mm thick, attached to larger valve near apex, becoming pendent, displayed against red inner valve surface. Croat 4834,11110.
Abundant, especially in the younger forest and along the shore; rare in the older forest. Flowers usually twice per year (sometimes three times per year), once in the early dry season (January to February, rarely as late as April), once in the early rainy season (usually July and August), and rarely again in the late rainy season. Fruit maturation is considerably more staggered, and mature fruits have been seen from February to August, often at the same time as flowering.
Though flowers are functionally unisexual, there are few conspicuous differences between staminate and pistillate flowers or inflorescences.
Known only from Panama, probably extending into both Costa Rica and Colombia. In Panama, known only on the Atlantic slope from tropical moist forest in the Canal Zone, Bocas del Toro, and San Bias, from tropical wet forest in Colon and Code, and from premontane wet forest in the Canal Zone and Colon.
See Fig. 302.
Protium tenuifolium Engler subsp. sessiliflorum (Rose) Porter, Ann. Missouri Bot. Gard. 56:475. 1969 P. sessiliflorum (Rose) Standl.; P. neglectum var. panamense Swart; P. neglectum var. sessiliflorum (Rose) Swart Anime, Chutras, Comida de mono
Functionally unisexual tree, to 18 m tall, to 27 cm dbh;
outer bark thin, flaking in firm sheets, lenticellate; inner bark moderately thick, reddish; sap slowly forming viscid, cloudy droplets after slash, with strong, sweet aroma;
69.
BURSERACEAE/TETRAGASTRIS499
stems glabrous, prominently brown-lenticellate. Leaves imparipinnate; petioles flattened above with sharp mar- gins; petiolules thickened at both ends; leaflets 5-9 (13), mostly oblong to oblong-elliptic, acuminate with blunt acumen, acute to rounded at base, often ± inequilateral, 10-20(27) cm long, 5-9(12) cm wide, glabrous except for minute papillae. Panicles axillary, near stem apex, the branches ferruginous; staminate inflorescences somewhat larger than pistillate (to 24 cm long); flowers 5-parted, sessile, 3.5-5 mm long; calyx to 2 mm long, cupulate, 5-lobed, minutely pubescent, the lobes acute; petals lanceolate, yellow-green, thickened medially on outside, ca 3 mm long, erect to somewhat spreading at anthesis;
stamens 10, alternately long and short, the longest to 2 mm long, the shortest to 1.3 mm long, scarcely longer than the prominent, pubescent disk; ovary ovoid, densely pubescent; style short, the ovary and style together ca 2.7 mm long; stigma 5-lobed, persisting in fruit. Capsules drupaceous, bearing 1-5 pyrenes, ovoid to depressed- globose, with 4 or 5 lobes (rarely 2- or 3-lobed), mostly 2-3 cm diam, red at maturity, usually rounded at both ends, the carpels 4 or 5, each dehiscing by a single valve, the valve falling free to expose the white mesocarp sur- rounding pyrene; pyrenes black, 1 per carpel, 1-seeded, ovoid in face view, irregular in side view. Croat 11109, 14822.
Common along the shore; occasional in the forest.
Flowers chiefly in April and May. The fruits mature chiefly in the middle of the rainy season (August to October).
Small bees, possibly Trigona, have been seen visiting the flowers. The fruits are edible.
Costa Rica and Panama. In Panama, known only from tropical moist forest in the Canal Zone, Chiriqui, Pa- nama, and Darien. Reported from tropical wet forest in Costa Rica (Holdridge et al, 1971).
TETRAGASTRIS Gaertn.
Tetragastris panamensis (Engler) O. Kuntze, Rev.
Gen. PI. 1:107. 1891 Anime
Dioecious tree, to 35 m tall and 60 cm dbh, scarcely buttressed; outer bark rough, unfissured, flaky. Leaves imparipinnate; petioles flattened on upper surface, nearly glabrous; leaflets 7-9 (11), ± oblong-elliptic, acuminate, acute to nearly rounded at base (lateral leaflets often inequilateral), 5-20 cm long, 2-7 cm wide, nearly gla- brous, stiff, the midrib arched. Panicles axillary (or ter- minal by abortion of stem apex), loosely branched, 2-15 cm long, with most exposed parts ± appressed- puberulent; flowers (4) 5-parted; pedicels ca 1 mm long;
calyx cup-shaped, shallowly lobed, to 1.5 mm long;
corolla yellowish-green, 3-5 mm long, lobed ca one- third its length, the lobes acute, thick, erect; stamens (8)10, included, abortive in pistillate flowers; disk annu- lar, (8)10-lobed; ovary ovoid, (4) 5-lobed and (4)5-locular, sparsely pubescent; style short, pyramidal; stigma 5-lobed.
Capsules drupaceous, purplish or reddish to brown, broadly turbinate to depressed-globose, round in cross
500 DICOTYLEDONEAE
section, to 2.5 cm long and 3.5 cm wide, usually with all 5 carpels developing, rarely 3 or 4, the thick wedge- shaped valves red within, falling free at maturity; meso- carp spongy, sweet; pyrenes 1 per carpel, 1-seeded, ca
1.5 cm long (including aril), attached subapically and situated between slender, red, platelike partitions of the main body, covered with a fleshy, sweet, white aril ca 2 mm thick. Croat 6823, 11195.
Common in the forest. Flowers chiefly in June and July (rarely as late as August and September); individuals of the species may flower every two years. The fruits mature mainly from March to the middle of May (rarely to July).
On BCI, fruits are generally found on the ground, though usually only the star-shaped main axis and loose valves remain. The seeds are quickly taken by large birds and mammals.
Belize to Peru and Brazil. In Panama, ecologically variable; characteristic of tropical moist forest (Tosi, 1971), known in the Canal Zone, San Bias, Veraguas, Panama, and Darien; known also from tropical dry forest in Code, from premontane moist forest in the Canal Zone and Panama, from premontane wet forest in Pa- nama and Darien, and from tropical wet forest in Colon.
Reported from premontane rain forest in Costa Rica (Holdridge et al., 1971).
See Fig. 303.
TRATTINNICKIA Willd.
Trattinnickia aspera (Standl.) Swart, Recueil Trav.
Bot. Neerl. 39:426. 1942 Protium asperum Standl.
Carano
Functionally dioecious tree, 25-50 m tall, to ca 75 cm dbh, weakly buttressed, the base becoming very rough- ened and warty; outer bark hard, dark brown; inner bark granular, tan; sap with faint, pleasant aroma (not charac- teristically burseraceous). Leaves deciduous, imparipin- nate; petioles flat above, with a raised marginal rib; rachis triangulate on upper surface, rounded below; petiole and rachis shortly pubescent; leaflets 7-11 (to 19 on juveniles), oblong-ovate to oblong-elliptic, acuminate, cuneate to cordate at base (sometimes inequilateral on lateral leaf- lets), 10-20(30) cm long, 4-6(12) cm wide, asperous and scabridulous on both surfaces, sparsely hispidulous below especially on veins, stiff, the margins entire and ± undu- late; lateral veins prominent below. Flowers 3-parted, functionally unisexual, in terminal panicles to 29 cm long, usually in dense glomerules, the branches sharply angulate, densely floriferous, hispidulous; pedicels to 4 mm long at anthesis, broadened apically, flattened; calyx sericeous inside, ca 2.7 mm long, caducous; corolla ± urceolate, ca 5 mm long, dull red (tinged with green in pistillate flowers), trilobate, the lobes acute, divided ca halfway to base in staminate flowers and one-fourth to three-fourths the way to base in pistillate flowers; stamens 6 (rarely 10), almost sessile (abortive in pistillate flowers);
filaments strap-shaped; anthers oblong, ca 2 mm long, introrse; ovary ± ovoid, glabrous, 2-2.7 mm long at
anthesis (reduced in staminate flowers); styles 2, short and thick. Fruits drupaceous, ovoid, ca 1 cm long, blunt or rounded at apex, smooth (drying ± wrinkled and acute at apex), indehiscent, violet-purple at maturity; pyrenes (1)2, each 1-seeded. Croat 11667, 11881, 13932.
Frequent in the old forest. Flowers in late July and August. The fruits mature during the dry season from January to April. Trees lose all their leaves during the early rainy season, but are renewed soon.
Easily distinguished from all other species by its sand- papery, compound leaves with the petiole flattened on the upper surface. Because of the differences in stamen size and condition of the ovary, the sex of any tree can be determined in the field by fallen flowers. Staminate flow- ers usually fall soon after the pollen has been shed, in most cases with the corolla still attached.
Fruits are probably consumed whole by large birds and mammals, since there is relatively little mesocarp.
Known only from Panama; characteristic of tropical wet forest, principally on the Atlantic slope (Tosi, 1971);
known also from tropical moist forest in the Canal Zone, Bocas del Toro, and Panama and from premontane wet forest in the Canal Zone (Pipeline Road).
See Fig. 304.