26 A ' Flora Neotropica FIG. 3. Stipule and colleter variation in the genera referred to the Condamineeae (sensu Hooker, 1873). A-C. <strong>Rustia</strong>. A. R. occidentalis (Delprete 5267, TEX), adaxial view. B. R. alba (Delprete & Verduga 6420, TEX), adaxial view. C. R.formosa (Kirkbride 5431, F), adaxial view. D. Kerianthera preclara (Prance et al. 24293, NY), adaxial view. E. <strong>Tresanthera</strong> condamineoides var. condamineoides (Steyermark & Rabe 96163, US), adaxial view. F-H. Condaminea corymbosa (Delprete & Webster 6104, TEX). F. View from above showing stipular insertion. G. Detail of base of stipule showing colleters in adaxial view. H. Habit of stipules. I, J. Pinckneya bracteata (Delprete & Sharif 6472, TEX). I. Adaxial view. J. Lateral view of stipule insertion (white area is petiole attachment). K, L. Pogonopus speciosus var. sandwithianus (Delprete & Apreza 6359, TEX). K. Adaxial view. L. Lateral view of stipule insertion and tuft of hairs in stipules' axils (white area is petiole attachment). M. Picardaea cubensis (Shafer 7850, NY), adaxial view. N. Kajewskiella trichantha (Kajewski 1667, A), adaxial view. O-Q. Chimarrhis. 0. C. cymosa (Webster et al. 9205, DAV), adaxial view. P. C. glabriflora (Delprete & Verduga 6423, TEX), abaxial view. Q. C. turbinata (Irwin et al. 55443, GH), adaxial view. R. Dioicodendron dioicum (Hammered et al. 5233, NY), adaxial view. S. Dolichodelphys chlorocrater (Ramirez & Cardenas 1017, COL), adaxial view. T. Molopanthera paniculata var. paniculata (Mori et al. 10328, NY), adaxial view. All drawn to the same scale, except G.
Introduction to the Neotropical Genera Studied 27 organs" (Robbrecht, 1988). Additional information Inflorescences about colleter structure and function can be found in The terminology and classification of inflores- Esau, 1965 (311); Halle, 1967 (95-97); Lersten, cences in the present work has been heavily influenced 1974a, 1974b, 1975; Robbrecht, 1988 (64-68); and by the recent work of Weberling (1992). More spe- Rogers, 1987. cifically, Weberling (1977) presented a comprehen- Colleters in the <strong>Rubiaceae</strong> are most commonly sive review of the generalized typology of <strong>Rubiaceae</strong> found inside of the stipules, usually at the basal por- inflorescences, and diagrammatic representations can tion, and are also frequently found inside the calyx, be found in Delprete (1996d: fig. 2B). When differat the base of the calyx lobes or alternate to them. ent interpretations on inflorescence morphology were For the group of genera under study, colleters have presented by various authors, Weberling's (1977, been consistently found inside the stipules at the basal 1992) conclusions were used. portion. Inflorescence position, in the genera studied, is When apical stipules are about to open, their in- most commonly terminal, with the exception of ternal colleters begin to secrete a resinous material Chimarrhis (axillary and subterminal). I consider the which is believed to protect the young vegetative buds axillary inflorescences of Chimarrhis to be the result against herbivory. The presence of"sticky" resinous of extreme reduction of the last terminal nodes exudates in vegetative buds has been observed in (Weberling, 1977, 1992; Robbrecht, 1988). <strong>Rustia</strong>, Condaminea, Pogonopus, and Chimarrhis. I <strong>Part</strong>icular attention should be called to the infloconsider absence or presence, arrangement, and gen- rescences ofPogonopus, Molopanthera, and Dioicoeral morphology of stipular colleters significant taxo- dendron, which have been variously described as ternomic characters, especially when used at the generic minal or terminal and lateral. Standley (1918, 1930a, and specific levels (Fig. 3). 1931 b, 1931 c) described the inflorescence of Pogon- In <strong>Rustia</strong>, stipular colleters are present in a trian- opus as "cymes arranged into terminal leafy panicles"; gular area on the base in R. occidentalis (Fig. 3A), in Steyermark (1974) as [literally translated from Spana single basal row in R. alba (Fig. 3B), or as a narrow ish] "densiflorous subcorymbose-cymose with basal area in R.formosa (Fig. 3C). In Picardaea (Fig. plurichothomous ramification"; Dwyer (1980) as "ter- 3M) stipular colleters are present in a single basal row. minal paniculate and with small lateral cymes," In <strong>Tresanthera</strong> (Fig. 3E) colleters are long-slender and though he described P. speciosus as having "inflorespresent on the base. In Condaminea (Fig. 3G) colleters cences terminal and axillary and thyrsoid-cymose"; are present in an irregular basal area on each of the Burger and Taylor (1993) [as in Dwyer, 1980] defour stipular units. In Pogonopus (Fig. 3K,L) the scribed Pogonopus with "inflorescences terminal, colleters are in one to two irregular basal rows and paniculate and subcorymbose-cymose," but P. intermixed with sericeous hairs. In Chimarrhis (Fig. speciosus with "inflorescences terminal or axillary." 30-Q) the position of the colleters is variable, but In my opinion (and according to Weberling, 1992), usually present on a basal area in C. cymosa (Fig. 30), Pogonopus, Molopanthera, and Dioicodendron have in a single basal row in C. glabriflora (Fig. 3P), or on terminal frondose paniculate inflorescences, with cya densely packed narrow basal area in C. turbinata mose branches subtended by foliose bracts, similar to (Fig. 3Q). In Dioicodendron (Fig. 3R) the stipular foliage leaves. The basal "leaves" (pherophylls) subcolleters are sparsely distributed in the basal area and tending the inflorescence branches are identical in size intermixed with golden densely pubescent hairs. In and shape to foliage leaves, and are gradually smaller Dolichodelphys (Fig. 1S) the colleters are sparsely toward the distal portion of the inflorescence. In gendistributed on the basal portion of the stipule. In eral, the presence of"leaves" (pherophylls) in termi- Molopanthera (Fig. 3T) only a few colleters are nal inflorescences of the <strong>Rubiaceae</strong> has led the above present at the base of the stipules and intermixed with cited workers to incorrectly describe the infloresgolden-sericeous hairs. [In Pinckneya (Fig. 3I,J) stipu- cences as lateral and cymose instead of terminal and lar colleters are sparsely distributed in a basal trian- paniculate. This argument was convincingly made by gular area. In Kerianthera (Fig. 3D) the colleters are Troll (1950) and again by Weberling (1977, 1992). sparsely distributed in an ovate area up to half the Inflorescence position of Chimarrhis is unique length of the stipules and (as in Pogonopus) are in- among the genera studied in having what I call "axiltermixed with gold-sericeous hairs. In Kajewskiella lary subterminal inflorescences," being borne in pairs (Fig. 3N) the colleters are long-slender, in a basal area in the leaf axils on the last terminal nodes. In my field one-third the length of the stipule and intermixed with observations of Amazonian species (i.e., C. glabrisericeous hairs]. flora, C. hookeri, C. barbata, and C. duckeana), the
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