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«29. 7. 2002 Phyton (Horn, Austria) Vol. 42 Fasc. 1 159-168 Typological Reinterpretation of the Inflorescences in Cariceae (Cyperaceae) By Abelardo C. ...»

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©Verlag Ferdinand Berger & Söhne Ges.m.b.H., Horn, Austria, download unter www.biologiezentrum.at

29. 7. 2002

Phyton (Horn, Austria) Vol. 42 Fasc. 1 159-168

Typological Reinterpretation of the Inflorescences in

Cariceae (Cyperaceae)

By

Abelardo C. VEGETTI*)

With 3 Figures

Received Jannuary 8, 2002

Key w o r d s : Cariceae, Cyperaceae. -Inflorescences, morphology, typology.

Summary

VEGETTI A. C. 2002. Typological reinterpretation of the inflorescences in Cariceae (Cyperaceae). -Phyton (Horn, Austria) 42 (1): 159-168, 3 figures. -English with German summary.

The aim of this work is to make a typological reinterpretation of Cariceae inflorescences, avoiding the particular analysis and terminology for specific taxa, since they would make the comparative studies more difficult. The inflorescence of the Cariceae is polytelic and truncated; it consists of a region of homogenized distal short paracladia (pseudoflorescence) and of a variable number of truncated long paracladia. In some species the pseudoflorescence is truncated and consequently the inflorescence is exclusively constituted of truncated long paracladia.

Zusammenfassung VEGETTI A. C. 2002. Typologische Neuinterpretation der Infloreszenzen der Cariceae {Cyperaceae). - Phyton (Horn, Austria) 42 (1): 159-168, 3 Abbildungen. Englisch mit deutscher Zusammenfassung.

Eine typologische Neuinterpretation der Canceae-Infloreszenzen wird vorgelegt, wobei die, für einzelne Taxa geprägte, spezifische Terminologie vermieden wird, da letztere den Vergleich erschwert. Die Infloreszenzen der Cariceae sind polytel und trunkat. Sie bestehen aus einer Region homogenisierter, distaler Kurzparakladien (Pseudofloreszenz) und aus einer variablen Anzahl trunkater Langparakladien. Bei manchen Arten ist auch die Pseudofloreszenz reduziert, in diesen Fällen besteht die Infloreszenz daher ausschließlich aus trunkaten Langparakladien.

*) Prof. Dr. Abelardo C. VEGETTI, Morfologia Vegetal, Facultad de Ciencias Agrarias (UNL), Kreder 2805, (3080) Esperanza, Provincia de Santa Fe, Argentina, e-mail: avegetti ©fca.unl.edu.ar ©Verlag Ferdinand Berger & Söhne Ges.m.b.H., Horn, Austria, download unter www.biologiezentrum.at Introduction The inflorescences of Cariceae exhibit complex ramification systems bearing female and male spikelets (Fig.1). Previous works by HAINES & LYE 1972, SMITH & FAULKNER 1976, KUKKONEN 1984, 1990, REZNICEK 1990 and TIMONEN 1985, 1989, 1993, 1998 show the differences in the structure of such inflorescences. The inflorescence morphology, the degree of closure of the peryginium, and the morphology of the rachilla are the key characters used to delimit the genera of Cariceae (YEN & OLMSTEAD 2000). According to TIMONEN 1993, there are different opinions in relation to the structure and phylogeny of sedge inflorescences. Thus, no general agreement has been reached for the classification of the genera and subgenera of the Cariceae tribe.

The typological interpretation of the inflorescence in Carex disticha Hudson and Kobresia Willd. has been performed by KUKKONEN 1984, 1990.

Unlike the inflorescences of dicotyledons and other monocotyledons, in those of Cariceae, the floral units (florescences) are not clear (TIMONEN 1985, 1989). Acording to this author (TIMONEN 1998), the monotelic and polytelic inflorescences are the fundamental types, but the inflorescences of the Cariceae should be considered as the third basic type, the caricoid inflorescence. In her opinion these inflorescences should be understood on the basis of their overall organization and not on the basis of small units, the spikelets. However, TROLL 1964 has generated the typological analysis to understand and to compare the angiospermous inflorescences that have similar patterns.

The aim of this work is to make a typological reinterpretation of Cariceae inflorescences, avoiding the particular analysis and terminology for specific taxa, since they would make the comparative studies more difficult.

Typological Considerations The Cyperaceae inflorescences typologically analyzed so far, are polytelic (MORA OSEJO 1960; KUKKONEN 1984, 1986; VEGETTI & TIVANO 1991;

VEGETTI 1992, 1994; HEINZEN & VEGETTI 1994; BROWING & GORDON-GRAY 1999).

The floral group in which the main axis and the branches of diverse order in a polytelic inflorescence ends, is called a florescence. The florescence at the end of the main axis is the main florescence, and the florescence at the end of each paracladium, is called a coflorescence (TROLL 1964; WEBERLING 1965, 1985, 1989).

Below the main florescence, branches with a structure similar to that of main axis can occur. These branches are known as paracladia. These paracladia can be either reduced only to the coflorescence, or second order paracladia can originate below it. In this way, the ramification of the system continues.

©Verlag Ferdinand Berger & Söhne Ges.m.b.H., Horn, Austria, download unter www.biologiezentrum.at

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Fig. 1. - A and B, Carex polysticha BOECK. - C and D, Carex rip aria CURT. var. chilensis (BRONGN. ex DUPERREY) KÜK.; C, Inflorescence; D, female pseudospikelet. - E, Carex sororia KUNTH. - References: fp, female pseudospikelet; mp, male pseudospikelet; p, pseudospikelet with male and female spikelets; fs, female spikelet.

In polytelic inflorescences, the main florescence may be developed or not. If the main florescence is not present, this means that truncation has taken place and the polytelic inflorescence is truncate. The truncation of the main florescence is generally synchronized with the homogenization of the distal portion of the main axis (WEBERLING & al. 1993). As a consequence, the distal portion of the short paracladia forms a pseudoflorescence (TROLL 1965).





©Verlag Ferdinand Berger & Söhne Ges.m.b.H., Horn, Austria, download unter www.biologiezentrum.at In Cyperaceae, paracladia diminishes in acropetal sequence and consequently, according to TIMONEN 1993, numeration of axes must be from base to apex (MOKA OSEJO 1960; MEERT & GOETGHEBEUR 1979; VEGETTI & TIVANO 1991; VEGETTI 1992, 1994; HEINZEN & VEGETTI 1994) rather than reverse numeration (KUKKONEN 1984, 1986, 1990; BROWNING & GORDON-GRAY 1999).

Inflorescence Typology in Cariceae DAHLGREN & al. 1985 consider that Caricoid-like spikelets are very reduced and the flowers are unisexual (Fig. 1 B, D). These reduced spikelets may come to be arranged in cones with their subtending bracts resembling glumes; so that these cones, though made of many spikelets, resemble simple spikelets (Fig. 1) and thus are called pseudospikelets (HAINE & LYE 1983; GOETGHEBEUR 1999).

In Cariceae, as well as in Cyperaceae, the florescences are spikelets.

These have undergone an important reduction: they are formed, as in the case of Carex L. female spikelet, by a single flower and the prophyll that wraps the spikelets (Fig. 1 B).

In the inflorescence of Carex disticha (Fig. 2 C), the main axis does not have a terminal spikelet. In other words: the main florescence is missing.

This species has a polytelic truncated inflorescence. Distally, on the main axis, there are axillary spikelets. These spikelets are distal paracladia reduced to a coflorescence (short paracladia, sPc). They are homogenized and altogether constitute a group, a pseudoflorescence (sHF). In Cyperaceae, this pseudoflorescence has been called pseudospikelet. It had been erroneously considered by KUKKONEN 1984 to be a main florescence (Fig. 2 B).

Therefore, this author states that Carex disticha doesn't present a simple florescence in the sense of EICHLER 1875/78, since pistillate flowers are not arranged on the main axis but on the branches (Fig. 2 B). In the pseudoflorescence flowers are always arranged on short paracladia (branches).

Consequently in his misinterpretation, KUKKONEN 1984 denominates the branches (paracladia) placed below the pseudoflorescence as coflorescences. Those branches (Fig. 2 C) are long paracladia without a coflorescence (truncated long paracladia). This misunderstanding led KUKKONEN to affirm that the structure of coflorescences is not uniform, since the number of staminate flowers diminishes in acropetal sequence (Fig. 2 B). Consequently, this author affirms that the main florescence and coflorescence are not composed of equivalent components. According to TROLL 1964 and WEBERLING 1989 the florescences (main florescence and coflorescence) are composed of equivalent components.

Truncation of the main florescence and homogenization of distal short paracladia, resulting in a pseudoflorescence, can also be seen in the inflorescences of various Cariceae species described by KUKKONEN 1990 (Fig. 2, ©Verlag Ferdinand Berger & Söhne Ges.m.b.H., Horn, Austria, download unter www.biologiezentrum.at

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Fig. 2. - Carex disticha HUDSON: A and B, the interpretation of Kukkonen 1984: Fig. 5;

C, the same as B, but with the interpretation of the present author. - References: HF, main florescence; Cofi_Cof36, coflorescences; sHF, pseudoflorescence; sPc, short paracladium; lPCi-lPcse, long paracladia.

©Verlag Ferdinand Berger & Söhne Ges.m.b.H., Horn, Austria, download unter www.biologiezentrum.at Fig. 3 A-C and Fig. 4 A and B) and TIMONEN 1985 (Fig. 4), 1989 (Fig. 2), 1993 (Fig. 1, Fig. 14 and Fig. 15). Remarkably, in all of these species as well as in Carex disticha, truncation of coflorescences and homogenization in long paracladia are also observed (Fig. 2 C).

In the inflorescences of species of Kobresia (KUKKONEN 1990, Fig. 2 A-D, Fig. 3 A-C, Fig. 4 A and B) the pseudoflorescence is reduced (Fig. 3 A). A further reduction is observed in Kobresia macrolepis MEINSH.

(KUKKONEN 1990) and K. pygmaea C. B. CLAEKE var. filiculmis KÜKENTH.

(KUKKONEN 1990) where the pseudoflorescence is truncated (Fig. 3 B). This means that in both species the inflorescence is exclusively constituted of truncated long paracladia (Fig. 3 B).

Thus we can draw the conclusion, that the inflorescence of the Cariceae (Fig. 2 C and Fig. 3 A and B) lacks a main florescence. It is a truncate polytelic inflorescence consisting of a region of homogenized distal short paracladia (pseudoflorescence) and of a variable number of long paracladia. The latter have no coflorescence (truncated long paracladia) and are composed of a variable number of homogenized secondary short paracladia. In some species the inflorescence consists of long paracladia only, as described for Kobresia macrolepis and K. pygmaea var. filiculmis (Fig. 3 B).

The variations in the structure of Cariceae inflorescences result from diversity in: 1) different sexuality in the flowers of the short paracladia;

2) the distribution of different types of short paracladia in the pseudoflorescence and in the long paracladia; and 3) the number and the degree of development of long paracladia. This determines different types of axes (TIMONEN 1993, 1998) and forms of inflorescences within the tribe (KUKKONEN 1990; REZNICEK 1990).

The inflorescences of Cariceae have undergone the following reductive processes: 1) A reduction in the development of florescences resulting in a single flower or a few flowered florescence; 2) the truncation of the main florescence and the coflorescences of long paracladia, and 3) a homogenization of short paracladia on the main axis and on long paracladia.

These reductive processes have determined the pattern of Cariceae inflorescences. Subsequently, reductive processes have modified the number and the degree of development of long paracladia. Thus, species with richly branched inflorescences have given rise to plants with small, less branched ones (KUKKONEN 1990; YEN & OLMSTEAD 2000).

Acknowledgements

The author thanks: I. KUKKONEN, the Scientific Editor and the Managing Editor of the Annales Botanici Fennici, who generously gave the permission to reproduce in this paper the Figure 5 from the work of KUKKONEN 1984 and the Fig. 3 A and the Fig. 3 D from KUKKONEN 1990; and F. WEBERLING for his critical review of the manuscript, and W. OBEEMAYER for the adaption of the scanned Fig. 2 and 3.

©Verlag Ferdinand Berger & Söhne Ges.m.b.H., Horn, Austria, download unter www.biologiezentrum.at

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References

BROWNING J. & GORDON-GRAY K. D. 1999. The inflorescence in southern African species of Bolboschoenus (Cyperaceae). - Ann. bot. fenn. 36: 81-97.

DAHLGREN R. M. T., CLIFFORD H. T. & YEO P. F. 1985. The Families of the Monocotyledons: Structure, Evolution and Taxonomy: VI-XII. Springer-Verlag: Berlin.

EICHLER A. W. 1875/78. Blüthendiagramme, 1-2. - Engelmann, Leipzig.

©Verlag Ferdinand Berger & Söhne Ges.m.b.H., Horn, Austria, download unter www.biologiezentrum.at GOETGHEBEUR P. 1998. Cyperacecie: 141-190. - In: KUBITZKIK., HUBER H., RUSALL P. J., STEVENS P. S. & STÜTZEL T. (eds.), The families and genera of vascular plants, 4. - Springer Verlag. Berlin.

HAINES R. W. & LYE K. A. 1972. Studies in African Cyperaceae VII. Panicle morphology and possible relationships in Sclerieae and Cariceae. - Bot. Not. 125:

331-343.

— 1983. The sedges and rushes of East Africa. - East African Nat. Hist. Soc.

Nairobi.

HEINZEN F. & VEGETTI A. 1994. Typology of the inflorescence in Cyperus corymbosus

var. subnodosus and C. rotundus (Cyperaceae). - Beitr. Biol. Pflanzen 68:

263-273.

KUKKONEN I. 1984. On the inflorescence structure in the family Cyperaceae. - Ann.

bot. fenn. 21: 257-264.

— 1986. Special features of the inflorescence structure in the family Cyperaceae.

- Ann. bot. fenn. 23: 107-119.

— 1990. The inflorescence structure of Kobresia myosuroides and related species of sect. Elyna (Cyperaceae). -Ann. bot. fenn. 27: 159-167.

MEERT M. & GOETGHEBEUR P. 1979. Comparative floral morphology of Bisboeckelereae and Cariceae (Cyperaceae) on the basis of the anthoid concept. - Bull.

Soc. Roy. Bot. Belg. 112: 128-143.

MORA OSEJO L. E. 1960. Beiträge zur Entwicklungsgeschichte und vergleichenden Morphologie der Cyperaceen. - Beitr. Biol. Pflanzen 35: 293-341.

REZNICEK A. A.. 1990. Evolution in sedges (Carex, Cyperaceae). - Canad. J. Bot. 68 (7):

1409-1432.

SMITH D. L. & FAULKNER J. S. 1976. The inflorescence of Carex and related genera. Bot. Rev. 42: 53-81.



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