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Anatomy and Morphology |
Department of Ecology, Evolution, and Marine Biology, University of California, Santa Barbara, California 93106-1467 USA; and Department of Plant Biology, Louisiana State University, Baton Rouge, Louisiana 70803 USA
Received for publication July 31, 2001. Accepted for publication December 13, 2001.
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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Key Words: Caesalpinioideae • Detarieae • Fabaceae • floral development • flower • Hymenostegia • Isoberlinia • Leguminosae • Microberlinia
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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; Breteler and Wieringa, 1999
; Wieringa, 1999
; Mackinder, 2000
), molecular systematics (Bruneau et al., 2000
, 2001
), comparative study of wood (Gasson, Trafford, and Matthews, 2000
), vegetative anatomy (Herendeen, 2000
), pollen (Banks and Klitgaard, 2000
; Banks, Klitgaard, and Crane, 2001
), reproductive biology (Lewis, Simpson, and Neff, 2000
), and comparative floral development (Tucker, 2000a
, b
, c
, d
, 2001a
, b
, 2002
).
Comparisons of floral development reveal the basis for the observed significant morphological distinctions among related taxa. These distinctions may be quantitative (e.g., heterochronic, or differences in timing of events) or qualitative (novel developmental events that are present in some groups and absent in others). While both types of distinctions occur in floral ontogeny of Detarieae, qualitative differences are of more interest because they have more evolutionary potential. The aims of this paper are (1) to compare floral ontogeny in representative taxa of the poorly known caesalpinioid legume tribe Detarieae sensu lato and (2) to use these data to seek correlations with groups currently considered to be clades (Bruneau et al., 2000
, 2001
; Mackinder, 2000
). The nine suprageneric "groups" within the tribe (Cowan and Polhill, 1981a
, b
; Polhill, 1994
) that were originally established by Léonard (1952
, 1957)
are not supported by molecular evidence (Bruneau et al., 2000
), but they provide a framework to assure representation of the major groups in the tribe. The Léonard groups represented here are Berlinia group (Isoberlinia, Microberlinia, Tessmannia), Hymenostegia group (Hymenostegia), and Detarium group (Phyllocarpus). Bruneau's work provides a rearranged set of relationships in Detarieae that is supported by Breteler (1995)
and Breteler and Wieringa (1999)
with minor differences that will be discussed later.
I have used an assemblage of floral developmental characters to group taxa in previous papers in this series. The first developmentally based group of taxa (previously treated) has a radially symmetrical flower, a circular floral apex, relatively small bracteoles directly after their initiation, all 21 floral organs, helical initiation of five sepals, and no ring meristem. Some taxa in this group have a medianly positioned first sepal (Amherstia, Brownea, and Tamarindus; Tucker, 2000c
), while other taxa in the group have the first sepal initiated nonmedianly (Crudia, Cynometra, Saraca, Schotia; Tucker 2000b
, 2001a
, b
). The genera Saraca and Crudia (Tucker, 2000b
, 2001a
) were treated in separate papers because their floral ontogeny diverges markedly in being apetalous but differing in the causal factors resulting in apetaly.
A second developmentally based group (having the "Omega" complex) contrasts sharply with the first in having a radially elongate and tangentially narrow floral apex, relatively large bracteoles directly after their initiation, and nonhelical sepal initiation. A ring meristem functions during floral organogeny in some taxa, and organ number at anthesis may be less than the 21 organs initiated. The first sepal initiated may be either median (Aphanocalyx, Monopetalanthus; Tucker, 2000a
) or nonmedian (Brachystegia; Tucker, 2000a
).
Floral symmetry of the Omega group of Detarieae may be essentially radial or actinomorphic (as in the present paper), or zygomorphic, based on the adaxial petal being larger or solitary and on suppression of some stamens (Tucker, 2000a
, 2002
). The petals are completely missing in Brachystegia, making assessment of symmetry more difficult. Two genera (Paraberlinia, Sindora) are in the Omega group but differ significantly from any other Detarieae studied and will be described together (unpublished data).
The Detarieae treated here initiate all 21 floral organs per flower, and flowers are essentially actinomorphic. There is little or no organ suppression in the taxa studied here, including species of Hymenostegia, Isoberlinia, Microberlinia, Phyllocarpus, and Tessmannia. Taxa in the same group but showing strong suppression of petals and/or stamens after their initiation are described in the second paper of this series (Tucker, 2002
).
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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Isoberlinia angolensis (Benth.) Hoyle & Brenan
Organography
The genus is in the Berlinia group of Léonard and includes 5–7 tree species, one in tropical Africa, the others in deciduous woodlands of Sudan and the Zambezi region. The medium-sized flowers of Isoberlinia angolensis (Fig. 1a, b) are in corymbose terminal panicles of racemes. The bracts are small and deciduous; bracteoles are large, covering the floral buds (Fig. 1a) and persistent after flowering. The flower (Fig. 1a, b, and e) is radially symmetrical with five subequal sepal lobes (upper two not fused) on a calyx tube, five subequal white petals (sepals and petals about the same length or petals slightly longer), ten fertile stamens with oblong pale yellow anthers and white filaments, and a stipitate ovary with elongate filiform style and a terminal stigma that is attached to the adaxial side of the hypanthium rim (Fig. 1d; Thompson, 1924
; Hutchinson, 1964
; Cowan and Polhill, 1981b
).
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Initiation of both stamen whorls is unidirectional (Fig. 12), beginning with the median abaxial and two laterals of the outer whorl concurrently. The last two outer-whorl stamens were just being initiated on the adaxial side in the same figure, and the first two inner-whorl stamen primordia were being initiated on the abaxial side (Fig. 12). Time of initiation overlaps in the two stamen whorls. All members of both stamen whorls are present in the midstage in Fig. 13. The carpel cleft is beginning in the same figure.
Isoberlinia angolensis, organ development
The bracteoles become thick and densely tomentose, forming a protective cover enclosing the floral bud (Figs. 1a and 17). The five sepal lobes elongate and overlap slightly, fused below into a calyx cup (Figs. 1c and 17). The two adaxial sepals remain discrete. The five petals begin to heighten and grow marginally as flat, ovate laminas (Fig. 21) at a height of 400–500 µm. Close to anthesis (Figs. 1b and 23) the petals appear subequal in height and shape. They become broad, sessile, and imbricate (Figs. 1a, 23, and 24).
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The carpel cleft becomes visible on the adaxial side at a height of 160–180 µm (Figs. 13–15). The cleft deepens as the margins expand (Fig. 16). By a height of 250 µm, the carpel is densely covered by trichomes (Figs. 19, 20, 22 and 23–25) that obscure further stages in development. At anthesis, the gynoecium is attached to the adaxial side of the hypanthium (Fig. 1d).
Microberlinia brazzavillensis A. Chev
Microberlinia, in the Berlinia group, includes two species of tall trees from tropical Africa with distichous branches and helical racemes or axillary panicles. The bracts are very large in bud, imbricate, covering the whole inflorescence, and soon becoming deciduous. The bracteoles are obovate, valvate, keeled, fused, and enclose the flower in bud (Fig. 26c and d). The radially symmetrical flower (Fig. 26a, b, and h) has four narrow uniform pale green sepals (Fig. 26f) that do not overlap at anthesis; the adaxial two are fused. The five subequal white petals include one that is clawed and slightly wider (Fig. 26g), up to 6.6 mm long and 3.2 mm wide, and four narrower petals, up to 2.2 mm wide but equally long or longer. The ten stamens include nine connate into a sheath split above (Fig. 26b), and the tenth is free; the anthers are yellow and uniform with white filaments. The ovary is pale green, stipitate, adnate to the margin of the tubular hypanthium (Fig. 26e). The style is slender and truncate; the pod valves have a strong wing (Hutchinson, 1964
; Cowan and Polhill, 1981b
; Wieringa, 1999
). Floral symmetry is essentially actinomorphic (Fig. 26h), except for the incomplete filament sheath open adaxially and the adaxial attachment of the gynoecium stipe.
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Petal initiation is unidirectional with the first pair abaxial and the second pair lateral (Figs. 33–35). The fifth, in median adaxial position, is difficult to discern until stamen initiation (Fig. 36). The carpel primordium initiates from the highly convex floral apex concurrently with the petals (Fig. 33).
Order of stamen initiation is bidirectional starting from the lateral positions in the outer antesepalous whorl and unidirectional from the abaxial side in the inner antepetalous whorl. The two whorls overlap with one other in time. The first stamen primordia of the outer antesepalous whorl to initiate are the two laterals (Figs. 33–36). The first two inner antepetalous stamens are also visible in these same figures. Next to initiate is the median abaxial outer stamen (Figs. 37 and 38). The remaining two antesepalous stamen primordia next initiate on the adaxial side; one is visible in Figs. 37 and 38; both are seen in Fig. 39. The lateral stamens of the inner antepetalous whorl initiate next (Fig. 39), and lastly the median adaxial of the same whorl is initiated, seen in the same figure. All five organ whorls overlap in time of initiation.
Microbarlinia brazzavillensis, organ development
The paired bracteoles are the protective structures, greatly overtopping the floral organs even in large buds (Figs. 26a and 48). The sepal primordia form a shallow, five-lobed calyx cup, with the abaxial sepal slightly larger than the rest, up to the time of midstage (Figs. 26b, f, and 41). Fusion between the two adaxial sepals is partial in Fig. 26f; the degree of fusion may vary, as Hutchinson (1964)
described the flower as four-sepallate, which would indicate complete fusion of these two. The sepals remain relatively thin and narrow (Figs. 26a, f, and 49); they overlap imbricately in bud but are separate at anthesis.
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). All ten stamens become functional. The outer-whorl antesepalous stamens go through differentiation before those of the inner antepetalous whorl. All stamen primordia of both whorls are equal in size at midstage (Figs. 40–42). The outer-whorl primordia become distally enlarged (Fig. 44) and broadened (Fig. 45). Anther and filament become evident (Fig. 47), and microsporangia begin to form, delimited by adaxial and lateral grooves (Figs. 49 and 50). The anthers are basifixed (Figs. 44 and 45) when first formed, but become dorsifixed later in development (Figs. 47, 49, and 54). Nine of the stamens become dorsifixed, connate, and raised up on a short filament tube (Figs. 26b and 49) split adaxially, formed by intercalary growth in the receptacle below the level of filament attachment. The median adaxial stamen remains free.
The carpel forms an adaxial cleft at a height of about 140 µm (Fig. 41). After elongating to a height of about 250 µm, ovule initiation is visible in the locule, seen between the gaping margins (Figs. 43 and 45). The carpellary margins remain open unusually long (Figs. 46 and 47) in this species. Eventually the carpellary margins become appressed and fuse. The gynoecium has a trichome-covered ovary, a revolute style in bud (Figs. 26a, 51, and 52), and a narrow stigma (Fig. 53). The carpellary stipe is attached to the adaxial side of the hypanthial rim (Fig. 26e).
Microberlinia bisulcata A. Chev
A complete organogenetic series was obtained for Microberlinia bisulcata, but only the early and midstages are illustrated because later stages do not differ appreciably from those in M. brazzavillensis. Bracts and subtended floral apices are initiated in helical acropetal order in the inflorescence (Fig. 56). Each floral apex is tangentially broad (Fig. 57) before bracteole initiation; the two bracteoles are initiated successively (Figs. 58 and 59) with broadly attached bases. Sepals are helical with the first median abaxial (Fig. 59) and the second median adaxial (Fig. 60). The last three sepals initiate in helical succession, but they remain as mere lateral shelves (at arrowheads in Figs. 61–63) through organogeny.
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Although few older stages are shown of M. bisulcata, the carpel cleft (at a carpel height of about 190 µm) and the calyx tube are beginning in Fig. 65. The lengthening calyx lobes are shown in Figs. 65 and 66. The carpel margins are still open at an approximate height of 400 µm at the time of ovule initiation (Fig. 67); they become sealed shortly thereafter (not shown).
Hymenostegia klainii Pierre ex Pellegr
Hymenostegia, in the Hymenostegia group of Cowan and Polhill (1981a)
, includes 16–20 species of trees and shrubs of tropical Africa. (This species is not illustrated, because its organogeny is very similar to that of Isoberlinia angolensis.) Hymenostegia klainii is a species of small trees about 3 m high with erect, racemose inflorescences that have pale pink helically arranged bracts. The bracteoles are large and ovate, white, petaloid, membranous, glabrous, flaring and persistent at anthesis. The two bracteoles fold imbricately in early stages but remain free and do not enclose the bud when large. The calyx in bud is pink. The small radially symmetrical flowers have a cupular or cylindric calyx tube, four imbricate sepals (the adaxial composed of two confluent sepals), five unequal free yellow petals (the outer two or three longer and broader, the abaxial two shorter and linear) that turn red over time. There are ten free stamens with white filaments and dorsifixed pale brown anthers that are inverted in bud and dehisce longitudinally. The ovary is pink with white hairs, stipitate, with filiform sigmoid style and punctiform white stigma. The carpel stipe is adnate to the side of the hypanthium (Hutchinson, 1964
; Cowan and Polhill, 1981b
; label information from F. J. Breteler).
Developmentally, Hymenostegia klainii has an Omega-type floral apex at bracteole initiation, massive bracteoles, helical initiation of five sepals (the first nonmedian and abaxial), and unidirectional initiation in the petal and stamen whorls. The carpel initiates after the petals. The stamen primordia remain essentially similar in size in each whorl during development.
Tessmannia africana Harms
Tessmannia includes approximately 12 tree species of tropical Africa and is in the Detarium group. (This species is not illustrated.) The species are large trees with terminal or axillary racemes or panicles of flowers. Tessmannia africana has short compact terminal panicles, each branch of which has about 8–15 flowers arranged distichously. Bracts of T. africana are small and caducous at about 1 mm in length; bracteoles are free and neither aestivate nor valvate. Floral buds are glutinous, covered with short brown peglike warts, and are 7–9 mm high before anthesis. Individual flowers of T. africana are radially symmetrical and have four greenish brown sepals, slightly imbricate or valvate, verrucose or strongly warty. The five subequal petals are pale pink, narrowly oblanceolate, broadly clawed with a plicate margin on the limb. The ten stamens are declinate in bud; nine are connate in a thick hairy sheath, and one is free. The filaments are pink, the anthers red and dorsifixed, dehiscing lengthwise. The ovary is stipitate and the style is long, circinately coiled, with a small capitate stigma. The genus (Cowan and Polhill, 1981b
) is characterized by radial symmetry, five subequal petals, ten stamens with nine connate at base, and a short hypanthium.
Developmentally, T. africana is similar to the species of Isoberlinia and Microberlinia illustrated. It has an Omega-type floral apex at bracteole initiation and massive, enclosing bracteoles at an early stage. Petals are initiated unidirectionally, and the carpel initiates with the petals. No other stages of organogenesis were available for T. africana. Late stages of floral development are obscured by an abundance of hairs covering most organs.
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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). The "Omega" assemblage includes the following character states: a laterally narrow, sagittally elongate post-bracteole floral apex that is tapered adaxially and abaxially, and bracteoles that are massive just after initiation that encircle 90% of the circumference of the post-bracteole floral apex. The bracteoles become large and enclose the floral buds. The taxa studied in the present paper (Isoberlinia angolensis, Microberlinia brazzavillensis, M. bisulcata, Hymenostegia klainii, and Tessmannia africana) have the "Omega" set of character states. Other character states that the taxa studied here share include radial floral symmetry, all 21 floral organs present, no floral organs suppressed, uniform calyx, uniform corolla, and all ten stamens alike and functional. Position of the first sepal varies; it is initiated medianly in Microberlinia and nonmedianly in Hymenostegia and Isoberlinia.
The character states that they share with the "circular apex" type group (Amherstia, Cynometra, Schotia, Tamarindus, Crudia, etc.; Tucker, 2000c
, 2001a
, b
) include helical initiation of sepals in most taxa and complete calyx and corolla initiated. These are probably plesiomorphic character states and are shared with other caesalpinioid tribes, particularly the tribe Caesalpinieae (Kantz, 1996
).
Comparisons among taxa studied
The developmental series shown (for Isoberlinia angolensis, Microberlinia brazzavillensis, M. bisulcata) are representative of the range of variations observed among taxa with all 21 floral organs initiated, radial symmetry, and little or no organ suppression. All share the Omega-shaped floral apex and massive bracteoles immediately after their initiation. All have helical initiation of sepals, starting abaxially. They differ in whether the first sepal initiates medianly (Microberlinia brazzavillensis, M. bisulcata) or nonmedianly (Isoberlinia angolensis, Hymenostegia klainii), and in petal order: helical (I. angolensis) or unidirectional (M. brazzavillensis, M. bisulcata, H. klainii). Stamens initiate in unidirectional order in each whorl in the species studied, except for bidirectional order in the outer whorl in M. brazzavillensis. An unusual feature in M. bisulcata is the ring meristem, on which the stamens are initiated. Overlap in time of organ initiation between whorls occurs in I. angolensis, M. brazzavillensis, and M. bisulcata but not in H. klainii. The carpel initiates concurrently with petals in all except H. klainii, in which it initiates slightly later, with the outer stamens. The carpel remains open through the time of ovule initiation in both species of Microberlinia, but not in the other species studied. Stamen differentiation is essentially alike in all, with the anthers basifixed at midstage and then becoming dorsifixed in late stage of development. Hypanthium formation is pronounced, with the perianth and stamens attached to the rim, in both species of Microberlinia and in H. klainii.
Ring meristem
The floral apex forms a ring meristem (a raised meristematic circular ridge that initiates petal and stamen primordia) in Microberlinia bisulcata but not in the other taxa examined. The ring meristem is a significant developmental innovation in other legume taxa, particularly papilionoid tribe Swartzieae (unpublished data) and the Brachystegia group of Detarieae (Tucker, 2000a
, d
). In Swartzieae, the ring meristem functions in greatly increasing the number of stamens per flower, but it produces only the basic ten stamens in M. bisulcata, Ateleia herbertsmithii (Tucker, 1990
), and the Brachystegia group of Detarieae (Tucker, 2000a
). I have speculated (Tucker, 2000d
) that the ring meristem could have arisen by accelerating "overlap" between whorls and by decreasing the time interval between initiations of organ whorls. Overlap between initiation of organ whorls occurs in I. angolensis, M. brazzavillensis, M. bisulcata, and Berlinia grandiflora, but not in H. klainii, among the taxa studied here. Overlap among organ whorls is found in some other detarioid legumes (e.g., Monopetalanthus durandii, Brachystegia boehmii, B. glaucescens, Saraca declinata, S. indica; Tucker, 2000a
, b
).
Specializations
At anthesis there are a few minor zygomorphic character traits in some taxa, such as adaxial attachment of the gynoecium in I. angolensis, M. brazzavillensis, and M. bisulcata, and there are differences in sinus depth between adjacent sepal lobes in several. Slight connation among stamen bases in M. brazzavillensis and M. bisulcata is another specialization. The persistent open carpel margins at ovule initiation in M. bisulcata is unusual; it is a heterochronic feature, but whether it is a specialization is open to question. Open carpels with ovules have been noted in 25 caesalpinioid species (Tucker and Kantz, 2001
); the majority are in tribes Caesalpinieae and Detarieae.
Systematic relationships
A comprehensive phylogenetic analysis based on both molecular and morphological data and including 71 of the 84 genera of tribes Detarieae and Macrolobieae has been published by Bruneau et al. (2000)
. Most of the taxa studied here belong to the Berlinia group of tribe Macrolobieae sensu Bruneau et al. (2000)
. The analysis by Bruneau et al. (2000)
however includes Tetraberlinia and Paraberlinia, both of which have marked organ suppression and zygomorphy and therefore will be published separately (Tucker, 2002
; unpublished data). In a phylogenetic analysis of Berlinia and other putatively related genera, Mackinder (2000)
found the genera Berlinia and Isoberlinia to be monophyletic and supported by morphology, but other clades remain cryptic and are not supported by known morphological states.
My investigations included only a few genera of the radially symmetrical members of Macrolobieae, so my comments are limited to those taxa. Floral development among these relatively unspecialized taxa, with their radial symmetry and little or no organ suppression, is essentially uniform. Shared developmental features of the Macrolobieae of Bruneau et al. (2000)
that have been investigated developmentally are (1) the "Omega"-type floral apex (narrow tangentially, wide in sagittal plane) during initiation of bracteoles and the early sepals; and (2) massive bracteoles at initiation with very wide basal attachment that quickly enclose the floral apex. Many Omega-complex taxa show reduced numbers of organs at petal and sepal initiation (Tucker, 2000a
) or failure of organs to develop after initiation (Tucker, 2001a
). In the species investigated here, however, there is no consequent diminution of organ primordia of sepals or petals, as there is in other detarioid taxa having the reduced floral apex characteristic of the Omega complex.
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FOOTNOTES |
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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Banks H. I. B. B. Klitgaard P. R. Crane 2001 Palynological contribution to the systematics of the Caesalpinioideae (Leguminosae). In Legumes down under, Abstracts, 7. The Fourth International Legume Conference, 2–6 July 2001. Australian National University, Canberra, Australia
Breteler F. J. 1995 The boundary between Amherstieae and Detarieae (Caesalpinioideae). In M. D. Crisp and J. J. Doyle [eds.], Advances in legume systematics, part 7, Phylogeny, 53–62. Royal Botanic Gardens, Kew, UK
Breteler F. J. J. J. Wieringa 1999 Generic delimitation in Caesalpinioideae. XVI International Botanical Congress, St. Louis, Missouri, August 1999. Abstract 16.2.1. Missouri Botanical Garden, St. Louis, Missouri, USA
Bruneau A. F. J. Breteler J. J. Wieringa G. Y. F. Gervais F. Forest 2000 Phylogenetic relationships in tribe Macrolobieae and Detarieae as inferred from chloroplast trnL intron sequences. In P. S. Herendeen and A. Bruneau [eds.], Advances in legume systematics, part 9, 121–149. Royal Botanic Gardens, Kew, UK
Bruneau A. F. Forest P. S. Herendeen B. B. Klitgaard G. P. Lewis 2001 Phylogenetic relationships in the Caesalpinioideae (Leguminosae) as inferred from chloroplast trnL intron sequences. Systematic Botany 26: 487-514[ISI]
Cowan R. S. R. M. Polhill 1981a Detarieae. In R. M. Polhill and P. H. Raven [eds.], Advances in legume systematics, part 1, 117–134. Royal Botanic Gardens, Kew, UK
Cowan R. S. R. M. Polhill 1981b Amherstieae. In R. M. Polhill and P. H. Raven [eds.], Advances in legume systematics, part 1, 135–142. Royal Botanic Gardens, Kew, UK
Gasson P. C. Trafford B. Matthews 2000 Wood anatomy of Caesalpinioideae. In Legumes down under, Abstracts, 30. The Fourth International Legume Conference, 2–6 July 2001. Australian National University, Canberra, Australia
Herendeen P. S. 2000 Structural evolution in the Caesalpinioideae (Leguminosae). In P. S. Herendeen and A. Bruneau [eds.], Advances in legume systematics, part 9, 45–64. Royal Botanic Gardens, Kew, UK
Hutchinson J. 1964 The genera of flowering plants (Angiospermae), vol. 1, Dicotyledones. Oxford University Press, Oxford, UK
Kantz K. E. 1996 Floral development in the caesalpinioid tribe Caesalpinieae (Fabaceae). Ph.D. dissertation, Louisiana State University, Baton Rouge, Louisiana, USA
Léonard J. 1952 Cynometrieae et Amherstieae. In Flore du Congo Belge et du Ruanda-Urundi, vol. 3, Spermatophytes, 279–495. Publications de l'Institut National pour l'Étude Agronomique du Congo Belge, Bruxelles, Belgium
Léonard J. 1957 Genera des Cynometreae et des Amherstieae africaines (Leguminosae-Caesalpinioideae). Memoires Academie Royale de Belgique, Classe Sciences 30: 1-314
Lewis G. P. B. B. Simpson J. L. Neff 2000 Progress in understanding the reproductive biology of the Caesalpinioideae (Leguminosae). In P. S. Herendeen and A. Bruneau [eds.], Advances in legume systematics, part 9, 65–78. Royal Botanic Gardens, Kew, UK
Mackinder B. 2000 The monophyly and taxonomic position of Berlinia Hook. f. (Leguminosae-Caesalpinioideae): a preliminary study. In P. S. Herendeen and A. Bruneau [eds.], Advances in legume systematics, part 9, 151–163. Royal Botanic Gardens, Kew, UK
Polhill R. M. 1994 Complete synopsis of legume genera. In F. A. Bisby, J. Buckingham, and J. B. Harborne [eds.], Phytochemical dictionary of the Leguminosae, vol. 1, xlix–liv. Chapman and Hall, New York, New York, USA
Thompson J. M. 1924 Studies in advancing sterility, part 1, The Amherstieae. Hartley Botanical Laboratory Publications, Liverpool University, Liverpool, UK
Tucker S. C. 1990 Loss of floral organs in Ateleia (Leguminosae: Papilionoideae: Sophoreae). American Journal of Botany 77: 750-761[CrossRef][ISI]
Tucker S. C. 2000a Evolutionary loss of sepals and/or petals in detarioid taxa Aphanocalyx, Brachystegia, and Monopetalanthus (Leguminosae: Caesalpinioideae). American Journal of Botany 87: 608-624
Tucker S. C. 2000b Floral development and homeosis in Saraca (Leguminosae: Caesalpinioideae: Detarieae). International Journal of Plant Sciences 161: 537-549[CrossRef]
Tucker S. C. 2000c Floral development in tribe Detarieae (Leguminosae: Caesalpinioideae): Amherstia, Brownea, Tamarindus. American Journal of Botany 87: 1385-1407
Tucker S. C. 2000d Organ loss in detarioid and other leguminous flowers, and the possibility of saltatory evolution. In P. S. Herendeen and A. Bruneau [eds.], Advances in legume systematics, part 9, 107–120. Royal Botanic Gardens, Kew, UK
Tucker S. C. 2001a The ontogenetic basis for missing petals in Crudia (Leguminosae: Caesalpinioideae: Detarieae). International Journal of Plant Sciences 162: 83-89[CrossRef][ISI]
Tucker S. C. 2001b Floral development in Schotia and Cynometra (Leguminosae: Caesalpinioideae: Detarieae). American Journal of Botany 88: 1164-1180
Tucker S. C. 2002 Comparative floral ontogeny in Detarieae (Leguminosae: Caesalpinioideae). 2. Zygomorphic taxa with petal and stamen suppression. American Journal of Botany 89: 888-907
Tucker S. C. K. E. Kantz 2001 Open carpels with ovules in Fabaceae. International Journal of Plant Sciences 162: 1065-1973[CrossRef]
Wieringa J. J. 1999 Monopetalanthus exit. A systematic study of Aphanocalyx, Bikinia, Icuria, Michelsonia and Tetraberlinia (Leguminosae, Caesalpinioideae). Wageningen Agricultural University Papers 99 (4): 1–320. Wageningen University, Wageningen, The Netherlands
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  M. A. Bello, J. A. Hawkins, P. J. Rudall, M. A. Bello, J. A. Hawkins, and P. J. Rudall Floral Morphology and Development in Quillajaceae and Surianaceae (Fabales), the Species-poor Relatives of Leguminosae and Polygalaceae Ann. Bot., June 1, 2008; 101(9): 1433 - 1434. [Abstract] [Full Text] [PDF]
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M. A. Bello, J. A. Hawkins, P. J. Rudall, M. A. Bello, J. A. Hawkins, and P. J. Rudall Floral Morphology and Development in Quillajaceae and Surianaceae (Fabales), the Species-poor Relatives of Leguminosae and Polygalaceae Ann. Bot., February 1, 2008; 101(3): 483 - 483. [Abstract] [Full Text] [PDF]
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 B. Marazzi and P. K. Endress Patterns and development of floral asymmetry in Senna (Leguminosae, Cassiinae) Am. J. Botany, January 1, 2008; 95(1): 22 - 40. [Abstract] [Full Text] [PDF]
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M. A. Bello, J. A. Hawkins, and P. J. Rudall Floral Morphology and Development in Quillajaceae and Surianaceae (Fabales), the Species-poor Relatives of Leguminosae and Polygalaceae Ann. Bot., December 1, 2007; 100(7): 1491 - 1505. [Abstract] [Full Text] [PDF]
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S. C. Tucker Floral ontogeny in Swartzia (Leguminosae: Papilionoideae: Swartzieae): distribution and role of the ring meristem Am. J. Botany, September 1, 2003; 90(9): 1271 - 1292. [Abstract] [Full Text] [PDF]
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 S. C. Tucker Floral Development in Legumes Plant Physiology, March 1, 2003; 131(3): 911 - 926. [Full Text] [PDF]
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S. C. Tucker Comparative floral ontogeny in Detarieae (Leguminosae: Caesalpinioideae). 2. Zygomorphic taxa with petal and stamen suppression Am. J. Botany, June 1, 2002; 89(6): 888 - 907. [Abstract] [Full Text] [PDF]
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