Difference between revisions of "Geonoma undata"

Taxonomic notes: - Geonoma undata is a member of a group of high elevation, Andean species (the G. undata clade, also including G. lehmannii, G. orbignyana, G. talamancana, and G. trigona). These species have been treated differently by both Wessels Boer (1968) and Henderson et al. (1995). They are closely related and three of them - G. lehmannii, G. orbignyana, and G. undata are difficult to distinguish from one another, and extremely complex internally. In fact, G. undata is the second most variable species of Geonoma. It differs from other species in this group by its prophyll surfaces which are ridged and densely tomentose with widely to closely spaced ridges, the ridges unequally wide, often dividing from and rejoining other ridges. (Henderson, A.J. 2011)/Palmweb.

Subspecific variation: - Five traits vary within this species (stem branching, stem type, leaf division, leaf plication, adaxial veins)(distal lip also varies but may be a result of hybridization, see under G. undata subsp. undata). Excluding stem branching and leaf division and the trait for which there are few data (stem type), the state distributions of the remaining two traits (leaf plication, adaxial veins) divide the specimens into three subgroups. One of these has non-raised adaxial veins and a discrete geographical range, and based on this is recognized as subspecies (subsp. stenothrysa). The second subgroup has plicate leaves and the third has nonplicate leaves. However, leaf plication is difficult to score in this species and does not divide the specimens into consistent subgroups. These specimens are therefore examined on a geographical basis. There are several geographically isolated subgroups, but usually too few specimens in each subgroup to test for differences. These subgroups are recognized as subspecies. There is an isolated subgroup in the Lesser Antilles (Dominica, Guadeloupe, and Martinique). There are only four specimens, but given their geographic isolation they are recognized as a subspecies (subsp. dussiana). There is an isolated subgroup in the Guayana Highland region of Venezuela and adjacent Brazil and Guyana. It differs from its nearest neighbors in Andean Venezuela in seven variables (stem height, number of pinnae, apical pinna width, apical pinna angle, prophyll length, rachilla length, rachilla width). It is recognized as a subspecies (subsp. appuniana).There is an isolated subgroup from the Tumuc-Humac mountains in French Guiana and Suriname. There are only two specimens, but the flower pits tend to be decussately arranged and there are few pinnae (3-4) compared with other specimens and they occur at lower elevations (600-620 m). These are recognized as a subspecies (subsp. tumucensis). There is an isolated subgroup from Central America in Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, and western Panama. It differs from the remaining specimens in seven variables (plant height, basal pinna width, apical pinna width, apical pinna angle, peduncle width, rachilla length, rachilla width). It is recognized as a subspecies (subsp. edulis). There is an isolated subgroup from Cerro Tacarcuna in Panama. There are only two specimens but they have distinctive, distantly spaced flower pits and are recognized as a subspecies (subsp. tacarcunensis). There is an isolated subgroup from Andean Venezuela in Carabobo state. The two specimens have distinctive, pinnate leaves with narrow, linear pinnae and are recognized as a subspecies (subsp. venezuelana). There is a subgroup from eastern Andean slopes in Ecuador that have distinctive, narrow, linear pinnae and are reported to be rheophytes. These are recognized as a subspecies (subsp. pulcherrima ). There is a subgroup from Andean Ecuador which has distinctive, narrow rachillae and distantly spaced flower pits. It differs from other Ecuadorian specimens in 17 variables (plant height, stem height, internode length, leaf number, rachis length, rachis width, number of pinnae, basal pinna length, basal pinna width, basal pinna angle, apical pinna length, prophyll length, peduncular bract length, interbract distance, peduncle width, rachilla length, rachilla width), and is recognized as a subspecies (subsp. skovii). The remaining specimens, from Andean regions of South America in Venezuela, Colombia, Ecuador, Peru, and Bolivia, are extremely variable and are not divisible into subspecies and are recognized as a single subspecies (subsp. undata). (Henderson, A.J. 2011)/Palmweb.

There are a total of ten subspecies;

1.) Geonoma undata subsp. appuniana, Brazil North, Guyana, and Venezuela. From 0°46-6°04'N and 59°50-66°04'W in the Guayana Highland region of Venezuela, Brazil, and Guyana at 1752 (810-2700) m elevation in montane rainforest. Leaves veins raised and rectangular in cross-section adaxially; basal pinna 3.8 (0.5-27.0) cm wide; apical pinna 6.4 (1.5-15.0) cm wide. Inflorescences prophyll margins with irregular, spine-like projections; flower pits usually spirally arranged, not distantly spaced. There is geographic discontinuity and specimens occur on many, isolated mountains, but there are too few specimens from each mountain to test for differences amongst them. There is geographical variation in this subspecies. Linear regression shows there are significant associations between elevation and three leaf and four inflorescence variables. Squared multiple R for the regression of rachis width on elevation is 0.40, basal pinna angle 0.29, apical pinna angle 0.58, prophyll length 0.23, rachilla width 0.23, fruit length 0.49, and fruit diameter 0.45. The rachis becomes thicker, pinna angles narrower, prophylls longer, rachillae thicker, and fruits longer and wider with increasing elevation. (Henderson, A.J. 2011)/Palmweb.

2.) Geonoma undata subsp. dussiana, Leeward Is., Windward Is. From 14°40-16°05'N and 61°00-61°40'W in Dominica, Guadeloupe and Martinique at medium elevations in lowland or montane rainforest. Leaves veins raised and rectangular in cross-section adaxially; basal and apical pinna width no data. Inflorescences prophyll margins with irregular, spine-like projections; flower pits usually spirally arranged, not distantly spaced. (Henderson, A.J. 2011)/Palmweb.

3.) Geonoma undata subsp. edulis, Costa Rica, Guatemala, Honduras, Mexico Southeast, Nicaragua, and Panamá. From 7°18-16°19'N and 80°06-93°15'W in Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, and Panama at 1523 (850-2400) m elevation in lowland or montane rainforest. Leaves veins raised and rectangular in cross-section adaxially; basal pinna 4.9 (0.8-13.7) cm wide; apical pinna 12.6 (5.7-23.2) cm wide. Inflorescences prophyll margins with irregular, spine-like projections; flower pits usually spirally arranged, not distantly spaced. There is geographic discontinuity and specimens occur in two areas separated by the lowlands of southern Nicaragua. There are few significant differences in variables between specimens from these two areas (basal pinna length, basal pinna width, prophyll length). There is variation in pit arrangement throughout the range of the subspecies, from spirally arranged to irregularly decussate or tricussate. A few specimens have the pits loosely spiraled proximally and tricussate or decussate distally, or tricussate proximally and decussate distally. One specimen (Evans 1459) from Honduras is larger in inflorescence size than other specimens from that area and has a narrow, elongate prophyll, more like that of subsp. hoffmanniana. The range of this subspecies overlaps with that of subsp. hoffmanniana in several places?northern Nicaragua, Atlantic and Pacific slopes of the Central Cordillera in Costa Rica, and both slopes on the Cordillera de Talamanca in Costa Rica and Panama. In this last area specimens are much larger in size than those from other areas (as are most specimens of subsp. hoffmanniana, which see). Hammel (2003) considered that specimens of subsp. edulis (as G. edulis) and larger, sympatric specimens of subsp. hoffmanniana (as G. hoffmanniana) were 'virtually indistinguishable' in this area. All specimens from the Cordillera de Talamanca and all others from Panama have much thicker peduncles than other specimens. (Henderson, A.J. 2011)/Palmweb.

4.) Geonoma undata subsp. pulcherrima, Ecuador. From 3°30-4°18'S and 78°30-78°41'W on eastern Andean slopes in southeastern Ecuador at 923 (750-1130) m elevation along river banks in areas subject to flooding. Plants are reported to be rheophytes. Leaves veins raised and rectangular in cross-section adaxially; basal pinna 0.7 (0.3-1.0) cm wide; apical pinna 0.8 (0.5-1.0) cm wide. Inflorescences prophyll margins without irregular, spine-like projections; flower pits usually spirally arranged, not distantly spaced. (Henderson, A.J. 2011)/Palmweb.

5.) Geonoma undata subsp. skovii, Ecuador. From 2°40-3°38'S and 78°00-78°30'W in Ecuador on eastern Andean slopes at 1778 (1470-1920) m elevation in montane rainforest. Leaves veins raised and rectangular in cross-section adaxially; basal pinna 1.3 (0.6-2.0) cm wide; apical pinna 7.3 (2.7-11.3) cm wide. Inflorescences prophyll margins with irregular, spine-like projections; flower pits usually spirally arranged, distantly spaced (Henderson, A.J. 2011)/Palmweb.

6.) Geonoma undata subsp. stenothyrsa, Colombia. From 6°01-6°54'N and 75°01-75°40'W in the Central Cordillera in Colombia (Antioquia) at 1675 (1500-1900) m in montane rainforest. One specimen (Bernal 190, neotype of G. euterpoidea) has leaves with more pinnae than the others (23 versus 8-13). Leaves veins raised and rectangular in cross-section adaxially; basal pinna 1.8 (0.7-3.8) cm wide; apical pinna 4.8 (0.1-8.5) cm wide. Inflorescences prophyll margins with irregular, spine-like projections; flower pits usually spirally arranged, not distantly spaced. (Henderson, A.J. 2011)/Palmweb.

7.) Geonoma undata subsp. tacarcunensis, At 8°09'N and 77°15'W on Cerro Tacarcuna on the Panama-Colombia border, at 1612 (1400-1825) m elevation in montane rainforest. Leaves veins raised and rectangular in cross-section adaxially; basal pinna 2.6 (2.4-2.7) cm wide; apical pinna no data. Inflorescences prophyll margins with irregular, spine-like projections; flower pits usually spirally arranged, distantly spaced. (Henderson, A.J. 2011)/Palmweb.

8.) Geonoma undata subsp. tumucensis, French Guiana, and Suriname. From 2°15-2°29'N and 54°25-54°45'W on the Tumuc-Humac mountains in French Guiana and Suriname at 610 (600-620) m in lowland rainforest. Leaves veins raised and rectangular in cross-section adaxially; basal pinna 3.0 cm wide; apical pinna no data. Inflorescences prophyll margins with irregular, spine-like projections; flower pits decussately arranged, not closely spaced. (Henderson, A.J. 2011)/Palmweb.

9.) Geonoma undata subsp. undata, Bolivia, Colombia, Ecuador, Peru, and Venezuela. From 10°56'N-17°48'S and 63°28-80°42'W in Andean regions of South America from Venezuela to Bolivia at 1964 (550-3370) m elevation in lowland or, more often, montane Rainforest. Leaves veins raised and rectangular in cross-section adaxially; basal pinna 3.0 (0.5-14.0) cm wide; apical pinna 10.6 (2.0-30.0) cm wide. Inflorescences prophyll margins with irregular, spine-like projections; flower pits usually spirally arranged, not distantly spaced. There is considerable variation in this widespread subspecies and there are many local morphotypes. These are difficult to distinguish from one another and are variable within themselves. In Venezuela, Colombia, Ecuador, Peru, and Bolivia there is a widespread morphotype (margaritoides morphotype). It has 25.4 (6.4-54.0) cm long and 4.3 (2.2-7.1) mm wide rachillae and 8.5 (5.5-12.5) mm long and 6.3 (4.2-8.9) mm wide fruits. This morphotype is relatively uniform throughout the northern part of its range, usually with elongate rachillae, often tricussately arranged flower pits, and small fruits (e.g., types of G. margaritoides and G. barthia). However, there are many exceptions, notably specimens with unusually short and thin, or unusually short and thick rachillae that occur in scattered localities. There is geographical variation in this morphotype. Linear regression shows there are significant associations between elevation and one plant, two leaf, and five inflorescence variables. Squared multiple R for the regression of plant height on elevation is 0.07, basal pinna angle 0.19, apical pinna angle 0.11, prophyll length 0.35, interbract distance 0.22, number of rachillae 0.76, rachilla width 0.05, and fruit diameter 0.19. With increasing elevation, pinnae have narrower angles. prophylls and interbract distances are longer, rachillae wider, and fruits larger. Some specimens from the Coastal Range in Venezuela (undata morphotype) occur at higher elevations and have greater interbract distances, longer peduncles, shorter rachillae, and longer and wider fruits. The type of G. undata has this kind of inflorescence. Some specimens from Venezuela (Andes and Peninsula Paraguaná, Falcón) and Colombia are distinctive in their shorter, thinner rachillae (densa morphotype). The type of G. densa has this kind of rachillae. Some specimens lack distal lips of the flower pits (megalospatha morphotype). These are hypothesized to be hybrids between the margaritoides morphotype and Geonoma trigona. They share with G. trigona the absence of an upper lip, and occur in two areas where G. trigona occurs (Ecuador, Peru). It is predicted that G. trigona will be found in Bolivia at the third hybrid locality. The type of G. megalospatha has this kind of flower pits. Specimens from the Western Cordillera of the Colombian Andes (iodolepis morphotype) have small leaves like those of G. orbignyana and inflorescences with short, thin rachillae 9.0 (7.0-11.0) cm long and 2.6 (2.1-3.5) mm wide. Fruits are 8.5 mm long and 6.3 mm wide. The type of G. iodolepis is of this morphotype. In the Venezuelan, Colombian, Ecuadorian, and Peruvian Andes, and also the Sierra Nevada de Santa Marta in Colombia, there is an extremely variable morphotype (weberbaueri morphotype). Specimens have plicate leaves, short, thick rachillae 19.0 (9.0-33.0) cm long and 5.7 (3.0-9.4) mm wide, and 12.5 (8.0-15.4) mm long and 8.7 (5.0-12.0) mm wide fruits. Most specimens with fruits have large-sized fruits, but a few specimens (Dodson 15213, Vásquez 26597) have small, globose fruits. This morphotype differs from the margaritoides morphotype in 10 variables. See under Geonoma orbignyana subsp. orbignyana for potential hybrids with that subspecies. There is geographical variation in this morphotype. Linear regression shows there are significant associations between elevation and three inflorescence variables. Squared multiple R for the regression of prophyll length 0.39, interbract distance 0.26, and peduncle length 0.33. Prophylls, interbract distances, and peduncles increase in length with increasing elevation. On eastern Andean slopes in Ecuador, particularly from Mera and Puyo, specimens (mera-puyo morphotype) have relatively slender inflorescences, 14.9 (9.8-21.0) cm long and 2.1 (1.5-2.5) mm wide rachillae, and 6.0 (5.9-6.0) mm long and 4.6 (4.1-5.0) mm wide fruits. Also on eastern Andean slopes in Ecuador, there are specimens (intermediate morphotype) which appear intermediate between the margaritoides morphotype and Geonoma orbignyana. They have small leaves and inflorescences, like the latter, but the bracts of the inflorescence are like those of the former. They have 16.8 (8.8-29.3) cm long and 3.8 (3.1-5.0) mm wide rachillae and 8.2 (7.6-8.8) mm long and 6.1 (5.8-6.3) mm wide fruits. One specimen (sira morphotype) from Peru (Huánuco) has small leaves like those of Geonoma orbignyana, slender rachillae 13.5 cm long and 1.9 mm wide, almost decussately arranged flower pits, and 8.9 mm long and 6.3 mm wide fruits. (Henderson, A.J. 2011)/Palmweb.

10.) Geonoma undata subsp. venezuelana. Venezuela. From 10°12-10°15'N and 68°07-68°10'W on the Coastal Range in Venezuela at 1362 (1275-1450) m elevation in montane rainforest. Leaves veins raised and rectangular in cross-section adaxially; basal pinna 0.5 (0.4-0.5) cm wide; apical pinna 1.7 cm wide. Inflorescences prophyll margins with irregular, spine-like projections; flower pits usually spirally arranged, not distantly spaced. (Henderson, A.J. 2011)/Palmweb.

Uses: Leaves are ocasionally used for packing material and thatch. The stem is used for posts. The black substance of mature fruits is used as a dye, also used to make baskets, hats, and other small hand-held items. (Henderson, A.J. 2011)/Palmweb.

G. dussiana is a synonym of undata -Palmnerd 01\10