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(Last Modified On 11/19/2012)
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(Last Modified On 11/19/2012)
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Family
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PIPERACEAE
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Contributor
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T. G. YUNCKER
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Description
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Herbs, shrubs, or small trees, not infrequently repent or scandent, often with nodose stems, glabrous or pubescent, vascular bundles scattered; leaves mostly alternate, sometimes opposite or verticillate, simple, usually entire, ranging in size from less than 1 cm. up to 30-40 cm. or more long, often with glandular or pellucid dots; stipules lacking or present and commonly more or less adnate to the petiole; flowers very small, usually green, perfect or unisexual, in racemose or more commonly spike-like inflorescences, these usually somewhat fleshy, pedunculate, terminal or leaf-opposed, sometimes axillary or multiple on a common peduncle; the flowers mostly densely crowded, each in the axil of a usually peltate or sub- peltate, sessile or stipitate, occasionally adnate or concave bract; perianth none; stamens usually 2-6, filaments mostly free, the anthers terminal, erect, 2-celled; ovary sessile or stipitate, 1-celled, 1-ovulate; fruit drupaceous, small.
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Note
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The Piperaceae, one of the larger families of flowering plants, is pantropic in its distribution. Species are especially abundant in Latin America. They extend from the Keys of Florida through the West Indies, Mexico, and Central America, south- ward to Chile and Argentina. They occur in greatest number throughout northern South America from Brazil to Peru and northward to Panama and Costa Rica. Some species are widespread in their occurrence but each country has a rather large number of endemics. The total number of valid American species is unknown but unquestionably amounts to many hundreds and may approximate 2,000 or more. A number of species of widespread distribution; e.g., Piper aduncurn, P. hispidum, P. marginatum, and others appear to exhibit an infinite capacity of varia- tion in minor characters of leaf size, shape, and indumentum. The extremes often appear quite distinct in themselves but study of large numbers of collections may reveal intergradations to such a degree that the extremes lose their seeming sig- nificance. The writer has been reluctant to maintain species or even varieties established by previous workers on such apparent variations when sufficient evi- dence indicates intergradations. It is probable that errors of judgment have been made but it is felt that such errors are on the side of conservatism. On this basis, a number of previously described species have been reduced to synonymy. At the same time, a number of novelties have been described as new to science. It is believed that with greater abundance of material, especially mass collections, and with correlated field study still further reductions may be in order. On the other hand, such studies may prove the validity of species which now appear to be the same. Small collections had been made in Panama previous to 1800 especially by Nee and by Haenke, and around the middle of the last century larger collections were prepared by Seemann, Fendler, Hayes, and others. However, collecting in Panama on an extensive basis did not get under way to any extent until after 1900. Miquel, the great Dutch authority and monographer of the Piperaceae during the first part of last century, studied some of the earlier collections. Casimir DeCandolle, third in line of the great Swiss family of botanists, following Miquel's death, published in the DeCandolle 'Prodromus' in 1869 a monograph of the Piper- aceae. Included in this publication were all of the species then known from Panama. Following this work, there appeared an almost continuous stream of papers by DeCandolle embodying the descriptions of hundreds of new species based on specimens sent to him by collectors as they explored farther and farther into botanically unknown areas. Following DeCandolle's death in 1918, Professor William Trelease, of the University of Illinois, undertook the study of the family, especially as it is represented in the Americas, and published a number of papers including the descriptions of many new species. The earliest paper to deal exclusively with the Piperaceae of Panama was pre- pared by DeCandolle and published posthumously in 1920. This included the descriptions of a number of new species based on collections made by Pittier and by Maxon in the early part of the century. Subsequently, large collections were made by Standley, Killip, Piper, and others. From a study of this rather considerable accumulation of material, Trelease in 1927 presented a monographic study of the Panamanian Piperaceae which included 139 species and a number of varieties. Woodson and his associates at the Missouri Botanical Garden have more recently made extensive collections. These have also been studied by Trelease who has pub- lished more than 70 new species and a number of varieties based on this material. Thus, according to Trelease, more than 200 species are represented in Panama. An attempt has been made in preparing the present paper to account for all species previously reported. Ninety-five species of Piper, 1 of Ottonia, 2 of Pothomorphe, 1 of Sarcorhachis, 1 of Anderssoniopiper, 72 of Peperomia, 22 varie- ties of Piper and 7 of Peperomia, making a total of 172 species and 29 varieties, are considered as valid in the present treatment. For the present study the writer has had the loan of the collections at Gray Herbarium of Harvard University, University of Illinois, Missouri Botanical Gar- den, New York Botanical Garden, and the United States National Museum. In addition, he has also had the loan of a number of critical specimens from Kew. He had the use of notes made by Trelease during his studies of type material in
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Note
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European herbaria.
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Key
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a. Stigmas 2-5 (mostly 3-4); floral bracts various, often more or less pubescent; plants more or less woody, commonly shrubby or vine-like, or less frequently herbaceous. b. Inflorescence leaf-opposed. c. Inflorescence spicate; fruit sessile .......................................................... 1. PIPER cc. Inflorescence racemose; fruit stipitate .................................................. 2. OTTONIA bb. Inflorescence axillary, spicate. c. Spikes several on a common peduncle .................................................. 3. POTHOMORPHE cc. Spikes one, or, if more than one, not on a common peduncle. d.Sikes solitary-4. SARCORHACHIS d. Spie soiay.....................................................4 S R O H C I dd. Spikes 2 (or more ?), superposed, axillary ...................................... 5. ANDERSSONIOPIPER aa. Stigmas 1 (may occasionally be cleft to simulate 2), stamens 2; floral bracts mostly round-peltate, glabrous; plants herbaceous, mostly com- paratively small ..................................... 6. PEPEROMIA
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