Morphological Phylogenetic Analysis of Two Early-Diverging Genera of Bromelioideae (Bromeliaceae)

Monteiro, Raquel Fernandes; Mantovani, André; Forzza, Rafaela Campostrini

doi:10.1590/2175-7860201566218

Abstract

Bromelia includes 66 species, grouped in three subgenera: Bromelia, Distiacanthus and Karatas. The genus is one of the earlier divergent groups within Bromelioideae, a subfamily with innumerable problems of generic delimitation. Considering that few phylogenetic studies have included more than one species of Bromelia, the objectives of this study were to carry out a phylogenetic analysis for the genus, seeking to evaluate its monophyly as well as its subgenera, and to better understand the interspecific evolutionary relationships within the genus and the intergeneric relationships within Bromelioideae. Also included in the analysis were the two species of Fernseea, a genus originally described from a species placed in Bromelia. The phylogenetic analysis was conducted with combined micro- and macromorphological data, including 39 taxa with 116 characters. As in other studies on Bromeliaceae, the clades have low statistical support and many questions regarding the relationships of the groups remain. The circumscription of Bromelia is still uncertain, due to the position of B. irwinii; Fernseea seems to emerge within this genus, as the marority-rule consensus suggests, and the infrageneric categories are debatable because they do not constitute monophyletic groups. However, Bromelia seems to be paraphyletic and, without B. irwinii, the genus can be considered monophyletic. Beginning with the premise that the most recent common ancestor of Bromelioideae appeared first in the southern Andes, Bromelia and Fernseea seems to have dispersed along three biogeographical routes. The present work is an important contribution to understand the evolution and biogeography of the Bromelioideae.

Key words:
Bromelia; Fernseea; dry diagonal; Brazilian Shield; morphology

[1] 2 Author for correspondence: raquelfmbio@yahoo.com.br

1	Leaf, rosette position in relation to central axis when sterile: 0, 15-30º;1, ca. 45º
2	Leaf, rosette position in relation to central axis during flowering: 0, 15-30º; 1, ca. 45º; 2, ca. 90º
3	Leaf, apex, orientation: 0, Erect; 1, Reflex
4	Leaf, apex, form: 0, Attenuate; 1, Acute; 2, Obtuse; 3, Emarginate; 4, Apiculate
5	Leaf, apex, pungent: 0, Present; 1, Absent
6	Leaf, margin: 0, Entire; 1, Aculeate
7	Leaf, spines, orientation: 0, One orientation, antrorse; 1, One orientation, retrorse; 2, Two orientations, antrorse and retrorse, on same leaf.
8	Leaf, spines, color: 0, Brown; 1, Green; 2, Red; 3, Yellow; 4, Wine-colored
9	Leaf, petiole: 0, Present; 1, Absent
10	Leaf, differentiation between sheath and blade: 0, Abrupt; 1, Tenuous
11	Leaf, blade, color when sterile: 0, Completely green; 1, Green to red at apex; 2, Completely red;
12	Leaf, blade, color of central leaves of rosette during blooming: 0, Completely green; 1, Green to red at base; 2, Completely red
13	Leaf, blade, consistency: 0, Carnose; 1, Chartaceous
14	Leaf, sheath, indumentum: 0, Present; 1, Absent
15	Leaf, sheath, indumentum position: 0, Only on adaxial face; 1, Only on abaxial face; 2, On both faces
16	Leaf, sheath, type of indumentum on adaxial face: 0, Lepidote; 1, Tomentose; 2, Floccose; 3, Villous; 4, Pannose; 5, Sericeous
17	Leaf, sheath, type of indumentum on abaxial face: 0, Lepidote; 1, Tomentose; 2, Floccose; 3, Villous; 4, Pannose; 5, Sericeous; 6, Strigose
18	Inflorescence, peduncle, size related to the rosette: 0, Inside, peduncle very reduced; 1, Emergent
19	Inflorescence, peduncle, indumentum: 0, Present; 1, Absent
20	Inflorescence, peduncle, type of indumentum: 0, Velutinous; 1, Tomentose; 2, Floccose; 3, Villous; 4, Lepidote; 5, Sericeous
21	Inflorescence, peduncle, cover by bracts: 0, Completely covered, imbricate bracts; 1, Partially covered, lax bracts
22	Inflorescence, peduncle, color: 0, Completely white; 1 Red; 2, Green; 3, Pink; 4, Brown
23	Inflorescence, bracts of peduncle, size related to the internode: 0, Longer; 1, Shorter
24	Inflorescence, bracts of peduncle, position related to the peduncle during anthesis: 0, Erect; 1, Patent, 45º-90º; 2, Completely reflexed
25	Inflorescence, bracts of peduncle, form: 0, Ovate; 1, Oblong; 2, Elliptical; 3, Lanceolate; 4, Linear
26	Inflorescence, bracts of peduncle, apex form: 0, Attenuate; 1, Acute; 2, Apiculate
27	Inflorescence, bracts of peduncle, apex pungent: 0, Present; 1, Absent
28	Inflorescence, bracts of peduncle, type of indumentum: 0, Lepidote; 1, Tomentose; 2, Floccose; 3, Villous
29	Inflorescence, bracts of peduncle, margin: 0, Entire; 1, Aculeate
30	Inflorescence, bracts of peduncle, color: 0, Stramineous with red apex; 1, Green; 2, Pink; 3, Wine-colored; 4, White; 5, Red
31	Inflorescence, type: 0, Raceme; 1, Capitulum, 2, Spike, 3, Corymb
32	Inflorescence, flowers, disposition: 0, Lax, exposing the branches; 1, Congested, covering the branches
33	Inflorescence, branches, development: 0, Highly developed, long; 1, Less developed, short
34	Inflorescence, viability of apex: 0, Sterile; 1, Fertile
35	Inflorescence, flowers, anthesis of flowers on each branch: 0, Opening upward; 1, Opening disorderly
36	Inflorescence, primary bracts, size in relation to branches of inflorescence: 0, Smaller; 1, Larger
37	Inflorescence, primary bracts, postion relative to peduncle during anthesis: 0, Erect; 1, Patent, 45º; 2, Completely reflexed
38	Inflorescence, primary bracts, form: 0, Ovate; 1, Oblong; 2, Elliptical; 3, Lanceolate
39	Inflorescence, primary bracts, apex form: 0, Attenuate; 1, Acute; 2, Obtuse
40	Inflorescence, primary bracts, apex pungent: 0, Present; 1, Absent
41	Inflorescence, primary bracts, type of indumentum: 0, Lepidote; 1, Tomentose; 2, Floccose;
42	Inflorescence, primary bracts, position of indumentum: 0, On apex only; 1, Covering primary bract; 2, On base only
43	Inflorescence, primary bracts, margin: 0, Entire; 1, Aculeate
44	Inflorescence, primary bracts, color: 0, Red; 1, Green; 2, Pink; 3, Stramineous
45	Inflorescence, floral bracts, size: 0, Shorter than ovary 1 Equaling or exceeding ovary, but not sepals, 2, Equaling or exceeding sepals, but not petals, 3, Equaling or exceeding petals
46	Inflorescence, floral bracts, color: 0, White; 1, Stramineous; 2, Red; 3, Pink; 4, Brown; 5, Blue; 6, Green
47	Inflorescence, floral bracts, form: 0, Ovate; 1, Oblanceolate; 2, Oblong; 3, Lanceolate; 4, Linear; 5, Triangular
48	Inflorescence, floral bracts, margin: 0, Entire; 1, Aculeate
49	Inflorescence, floral bracts, carin: 0, Present; 1, Absent
50	Inflorescence, floral bracts, indumentum: 0, Present; 1, Absent
51	Inflorescence, floral bracts, type of indumentum: 0, Lepidote; 1, Tomentose; 2, Floccose, 3, Villous
52	Inflorescence, floral bracts, position of indumentum: 0, On apex only; 1, Covering floral bract
53	Inflorescence, floral bracts, apex: 0, Acute; 1, Attenuate; 2, Obtuse; 3, Rounded
54	Flower, sepal, form: 0, Ovate; 1, Obovate; 2, Oblong; 3, Lanceolate; 4, Linear; 5, Triangular; 6, Elliptical
55	Flower, sepal, symmetry: 0, Symmetrical; 1, Asymmetrical
56	Flower, sepal, color: 0, White; 1, Brown; 2, Blue; 3, Green; 4, Pink; 5, Stramineous
57	Flower, sepal, indumentum: 0, Present; 1, Absent
58	Flower, sepal, type of indumentum: 0, Lepidote; 1, Tomentose; 2, Floccose; 3, Villous
59	Flower, sepal, position of indumentum: 0, On apex only; 1, Covering sepal; 2, On base only
60	Flower, sepal, apex: 0, Acute; 1, Attenuate; 2, Obtuse; 3, Rounded; 4, Retuse; 5, Apiculate
61	Flower, sepal, consistency: 0, Membranaceous; 1, Carthaceous; 2, Coriaceous
62	Flower, sepal, carena: 0, Present; 1, Absent
63	Flower, sepal, margin: 0, Entire; 1, Aculeate; 2, Crenate
64	Flower, sepal, concrescence: 0, Free; 1, Conate
65	Flower, petal, position of apex during anthesis: 0, Cuculate; 1, Erect; 2, Recurvate
66	Flower, petal, consistency: 0, Membranaceous; 1, Carnose
67	Flower, petal, color of blade: 0, White; 1, Pink; 2, Red; 3, Lilac; 4, Purple; 5, Wine-colored; 6, Green; 7, Blue; 8, Black; 9, Yellow
68	Flower, petal, color of margin: 0, The same as the blade; 1, Different from the blade
69	Flower, petal, indumentum: 0, Present; 1, Absent
70	Flower, petal, apex: 0, Acute; 1, Attenuate; 2, Obtuse; 3, Rounded
71	Flower, petal, form: 0, Oblanceolate; 1, Oblong; 2, Obovate; 3, Elliptical; 4, Linear; 5, Triangular
72	Flower, petal, concrescence: 0, Free; 1, Conate
73	Flower, petal, kind of concrescence: 0, Up to1/3; 1, More than 1/3
74	Flower, petal, appendix: 0, Present; 1, Absent
75	Flower, petal, callosity: 0, Present; 1, Absent
76	Flower, stamens, length related to the petals: 0, Exceeding; 1, Not exceeding
77	Flower, stamens, position related to the petals: 0, Exserted; 1, Included
78	Flower, stamens, color of anthers: 0, White; 1, Yellow
79	Flower, stamens, concrescence in relation to petals: 0, Adnate; 1, Free
80	Flower, stamens, anthers insertion: 0, Basifixed; 1, Dorsifixed
81	Flower, stamens, anthers form: 0, Oblong; 1, Sagittate; 2, Elliptical
82	Flower, ovary, position: 0, Superior; 1, Inferior
83	Flower, ovary, form: 0, Ovate; 1, Elliptical; 2, Oblong; 3, Obovate
84	Flower, hypanthium, indumentum: 0, Present; 1, Absent
85	Flower, hypanthium, type of indumentum: 0, Villous; 1, Tomentose; 2, Floccose
86	Flower, hypanthium, sulcus: 0, Present; 1, Absent
87	Fruit, type: 0, Berry; 1, Capsule
88	Fruit, concrescence of external walls: 0, Free; 1, Concrescent
89	Fruit, petals: 0, Persistent; 1, Not persistent
90	Fruit, color: 0, Brown; 1, Green; 2, Orange; 3, Yellow; 4, Wine-colored; 5, Pink; 6, Red; 7, White
91	Seeds, appendix: 0, Present; 1, Absent
92	Leaf anatomy, abaxial surface, contour: 0, Smooth to slightly wavy; 1, Grooved
93	Leaf anatomy, epidermal cells of adaxial surface, thickening type: 0, Strongly thickened inner periclinal wall, reduced lumen; 1, Inner periclinal wall thickened, with a smooth, wide lumen; 2, Strongly thickened outer periclinal wall, reduced lumen
94	Leaf anatomy, epidermal cells of abaxial surface, thickening type: 0, Strongly thickened inner periclinal wall, reduced lumen; 1, Inner periclinal wall thickened, with a smooth, wide lumen; 2, Strongly thickened outer periclinal wall, reduced lumen
95	Leaf anatomy, trichome, number of pedicle cells: 0, Two; 1, More than two
96	Leaf anatomy, trichome on adaxial surface: 0, Present; 1, Absent
97	Leaf anatomy, stomatal apparatus, position related to epidermis: 0, Same level; 1, Down, 2, Up
98	Leaf anatomy, mechanical hypodermis adaxial, thickening: 0, Present; 1, Absent
99	Leaf anatomy, mechanical hypodermis adaxial, number of layers: 0, One; 1, Two; 2, Three; 3, Four; 4, Five
100	Leaf anatomy, mechanical hypodermis adaxial, thickening of cell: 0, Slightly thickened; 1, Strongly thickened
101	Leaf anatomy, mechanical hypodermis abaxial, number of layers: 0, One; 1, Two; 2, Three; 3, Four; 4, Five
102	Leaf anatomy, mechanical hypodermis abaxial, thickening of cell: 0, Slightly thickened; 1, Strongly thickened
103	Leaf anatomy, water-storage hypodermis adaxial, size: 0, Up to 1/3 of the blade thickness; 1, More than 1/3 of the blade thickness
104	Leaf anatomy, transition between adaxial water-storage hypodermis and chlorenchyma: 0, Abrupt; 1 Tenuous
105	Leaf anatomy, adaxial water-storage hypodermis cell shape: 0, Rounded, 1, Elongated in anticlinal direction
106	Leaf anatomy, water-storage hypodermis, position: 0, Adaxial face; 1, Abaxial face; 2, Both sides
107	Leaf anatomy, brachiform parenchyma, cell shape: 0, With short arms (forming small gaps); 1, With long arms (forming large gaps)
108	Leaf anatomy, cell of bundle sheath parenchyma, disposition: 0, Radiate; 1, Not radiate
109	Leaf anatomy, vascular bundles of smaller diameter, shape of the thickening (fiber): 0, Taller than wide; 1, Wider than tall
110	Leaf anatomy, vascular bundles of larger caliber, thickening of fibers: 0, Calotte; 1, Circling totally
111	Leaf anatomy, palisade chlorenchyma, shape: 0, Palisade; 1, Not palisade; 2, Palisade and not palisade
112	Leaf anatomy, adaxial epidermis, outer periclinal walls: 0, Smooth; 1, Papillose
113	Leaf anatomy, raphides: 0, Present; 1, Absent
114	Leaf anatomy, secretory ducts: 0, Present; 1, Absent
115	Leaf anatomy, extravascular fibers: 0, Present; 1, Absent
116	Habit: 0, Terrestrial; 1, Rupiculous; 2, Epiphytic

Brasil

Brasil

Morphological Phylogenetic Analysis of Two Early-Diverging Genera of Bromelioideae (Bromeliaceae)

Abstract