| BEN |
| BOTANICAL ELECTRONIC NEWS |
|---|
| ISSN 1188-603X |
| No. 406 March 26, 2009 | aceska@telus.net | Victoria, B.C. |
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Carroll Emory Wood, Jr., Harvard University Professor of Biology and Curator of the Arnold Arboretum, died March 15th at his home. He was teacher and mentor to many botanists and students here, at the University of North Carolina and at the University of Michigan Biological Station. A specialist in the flora of the Southeastern U.S., he initiated, supervised and edited a comprehensive flora of that region, “The Generic Flora of the Southeastern United States.” His fieldwork took him throughout eastern North America, the Caribbean and central Mexico.
He was admired by his many friends and colleagues for his compassion, commitment and unfailing sense of humour. A resident of Boston’s South End for almost 50 years, he was beloved by neighbours and friends for his gentle nature and horticultural guidance. A memorial “Celebration of Carroll” will be held Sunday, April 26, 2009, from 3-5 pm, at United South End Settlements, 566 Columbus Avenue, Boston.
The family Passifloraceae is not covered in any of the published manuals on Californian flora. However, four species of Passiflora (passionflowers) have been recently recognized as naturalized or, at least, casual in this state. We describe the largest infestation of Passiflora tarminiana Coppens & Barney, comment on all other reports of naturalized Passiflora taxa in California, and present a key for their identification. Passiflora tarminiana (until recently confused with P. mollissima Kunth in H.B.K.) and P. caerulea L. can be classified as naturalized and invasive, P. mixta L. as naturalized, P. manicata (Juss.) Pers. as a casual alien, and Passiflora ‘Coral Sea’ as persisting after cultivation in California.
Supplementary data at URL: http://bomi.ou.edu/ben/406/passiflora_figures.pdf
More than 530 species, mostly vines or lianas, have been described in the genus Passiflora (passion flowers) (Hansen et al. 2006), with many taxa popular in cultivation (Ohle 1975, Vanderplank 2000, Ulmer and MacDougal 2004). At least six of them are among the most aggressively invading weeds in several countries (Heenan and Sykes 2003, Randall 2002, Weber 2003). There are no native species in this genus in California. The whole family, Passifloraceae, is essentially tropical. The native distribution of the genus Passiflora in the western North America reaches its northern limits in Arizona and Baja California (Goldman 2003). The family Passifloraceae is not covered in The Jepson Manual (Hickman 1993). However, even if the Californian climate is, in general, not suitable for most species in this genus, more than ten different taxa are commonly cultivated in Californian gardens. Moreover, four species of Passiflora have been recently recognized as naturalized or at least as casual aliens (for terminology see Pysek et al. 2004) in California (Hrusa et al. 2002). Consequently, this genus deserves more attention. Our recent discovery of a large, spreading population of Passiflora tarminiana Coppens & Barney in Julia Pfeiffer Burns State Park, Monterey County, became an impetus for writing this note.
Here we comment on the four naturalized/casual Passiflora species in California, describe the new and largest infestation of P. tarminiana, and provide a key for identification of discussed taxa.
Traditionally, the genus Passiflora has been subdivided into 22 to 24 subgenera (Escobar 1989, Vanderplank 2000). However, most of them now correspond to supersections, sections, or series in Ulmer and MacDougal (2004).
Passiflora caerulea L. (section Granadillastrum) This South American species was reported from several localities in North Coast Ranges, San Gabriel Mountains, and South Coast by Hrusa et al. (2002). Passiflora caerulea is now widely naturalized in southern California (A. Sanders, personal communication). It is also naturalized and weedy in New Zealand (Webb et al. 1988), Australia (Lazarides et al. 1997), and South Africa (Henderson 2001).
Passiflora manicata (Juss.) Pers. (section Manicata) This species is native to the Andes of South America. There is one record of Passiflora manicata from Santa Barbara Co. (Smith 1976), although, it was not mentioned in the 2nd edition (Smith 1998). This species is reported as 'sparingly naturalized' on Kaua'i, Hawai'i (Wagner et al. 1999).
Passiflora ‘Coral Sea’ (Figs. 1 & 2) This is a hybrid between Passiflora manicata and an unknown species or several species (Ulmer and MacDougal 2004). While reported only recently (Dean et al. 2008), it is commonly cultivated along Californian coast. Extensively branching individuals can cover large trees (Figs. 2). Plants are sterile, but persisting for a long time after cultivation. Photograph of this taxon, under the name P. jamesonii (Mast.) Bailey, is on p. 69 in Phillips & Rix (1997).
Passiflora mixta L. (section Tacsonia) A local patch of this vigorous South American vine was reported by Hrusa et al. (2002) from Golden Gate Park. This is a relatively rare weed in New Zealand (Heenan & Sykes 2003).
Passiflora tarminiana Coppens & Barney (section Elkea; Figs. 3 & 4) In California, it has been so far found as established in disturbed habitats in Contra Costa Co. and reported from Riverside Co. (Hrusa et al. 2002). While existence of this taxon was anticipated earlier (de Wilde 1975, p.15), it was formally described as a distinct species only in 2001 (Coppens d’Eeckenbrugge et al. 2001). Passiflora tarminiana is not known from natural habitats, but it is cultivated and spreading in disturbed habitats in Venezuela, Colombia, Ecuador, and Peru. Presumably, it is native somewhere in this range. This is the famous 'banana poka', a serious pest in Hawaii where it has been called P. tripartita (Juss.) Poir. var. mollissima (Kunth) Holm-Nielsen & Jřrgensen [syn.: Passiflora mollissima Kunth in H.B.K.] until recently (Ramadan et al. 2008). Confusion with the previous species was unavoidable and pictures of P. tarminiana were often presented as those of P. mollissima (e.g., Henderson 2001, Muyt 2001, Vanderplank 2000, Wagner et al. 1999). Probably most of the material that is provided by nurseries in California under 'P. mollissima' is Passiflora tarminiana. Banana poka was classified as one of the twelve most significant plant invaders in Hawaii (Smith 1985). It topples native vegetation and suppresses tree regeneration, while its prolific fruit production leads to an increase of feral pig population densities. Its reproduction combines outcrossing early in anthesis with selfing at a later stage, ensuring that isolated individuals can produce viable offspring from a single colonizer (La Rosa 1992). Germination seems to be slow - Williams and Buxton (1995) reported that only 25% of fresh seeds germinated over the first 12 weeks, therafter germination slowed considerably. Vegetative reproduction was reported as rare in Hawaii (La Rosa 1992) but was found important at some sites in New Zealand where P. tarminiana is one of the most common invasive passion flowers (Heenan & Sykes 2003, Williams and Buxton 1995). Passiflora tarminiana is also naturalized in Australia and Papua New Guinea (Australia's Virtual Hebarium - http://www.chah.gov.au/avh ). Based on their description (de Wide 1975), it seems that about half of the specimens of ‘P. mollissima' from Kenya are also P. tarminiana.
In November, 2003, we found that Passiflora tarminiana is spreading around McWay Creek, by the entrance to Julia Pfeiffer Burns State Park, 25 km SE of Point Sur, Monterey Co. (360 09' 32'' N, 1210 40' 11'' W, elev. 50- 60 m). In total, seven distinct patches could be recognized at that time. Passiflora tarminiana was climbing up to 12 m on trees, including Alnus rhombifolia Nutt., Sequoia sempervirens (Lamb.ex D. Don) Endl., and Umbellularia californica (Hook. & Arn.) Nutt.(Fig. 4). We estimated that the extent of gross infestation was ca. 1.1 ha and net infestation ca. 0.1 ha. Consequently, this species can be now classified not only as 'naturalized' but also as 'invasive' (sensu Pysek et al. 2004) in California. We estimated that there were over 3000 fruits present in the canopy and on the ground in the infested area. Based on five randomly collected fruits, we estimated that the mean number of viable seeds per fruit is around 170 (142-204). The shape of seeds is quite variable and some of them match better the "P. mollissima" photograph in Pérez-Cortéz et al. (2002) than the drawing in the original description of P. tarminiana by Coppens d’Eeckenbrugge et al. (2001). At least some seeds are not cordate but rather ovate. A voucher specimen and photographs were deposited at DAV.
Depending on availability of dispersal agents (La Rosa 1984, Jacobi and Warshauer 1992, Williams et al. 2000), spread of this species can be quite fast. With feral pigs common in Monterey Co. (Pine & Gerdes 1973, Long 2003), they could become the major dispersers of Passiflora tarminiana in this area. Nevertheless, eradication of an infestation of this size, in general, is feasible (Rejmánek and Pitcairn 2002) and P. tarminiana populations of this size were successfully eradicated in New Zealand (P. A. Williams, personal communication). Based on our observations in recent years (2006, 2008), it seems that attempts to eradicate this species in Julia Pfeiffer Burns State Park have been under way. This is a beautiful ornamental plant, however, the only long-term profit from its spread could be suppression of Cape ivy (Delairea odorata Lem.), another invader, that is also present in the same area.
The following key to casual and naturalized Passiflora taxa in California (in bold) was compiled on the basis of Coppens d’Eeckenbrugge et al. (2001), Heenan and Sykes (2003), Holm-Nielsen (1988), Vanderplank (2000), Ulmer and MacDougal (2004), and examination of herbarium specimens.
Passiflora tripartita is included in the key as it is commonly confused with P. tarminiana (see above). It is a South American species, probably only occasionally cultivated in California. Hypanthium/sepal length ratio and stipule size and their persistence are important characters separating P. tarminiana and P. tripartita. For a long time, P. tripartita var. mollissima was classified as a separate species - P. mollissima . However, this taxon has been shown, using morphological and molecular data, to be a part of the natural variation of P. tripartita (references in Heenan & Sykes 2003). On the other hand, plants now classified as P. tripartita var. azuayensis Holm-Nielsen & Jřrgensen were previously often confused with P. mixta. Both varieties are common invaders in New Zealand (Heenan & Sykes 2003). P. mixta, P. tarminiana, and P. tripartita are closely related (Segura et al. 2002) and their hybrids are sometimes reported.
Judging from their persistence after cultivation, two other commonly planted Passiflora species are candidates for naturalization in California: P. incarnata L. (series Passiflora, native to eastern U.S.) and P. vitifolia Kunth (supersection Coccinea, native from Central America to Peru). Passiflora laurifolia L. (supersection Laurifolia, native in West Indies and Venezuela), occasionally cultivated in California, is naturalized in Hawaii and several other Pacific islands. The last three species are not included in this key, but their descriptions and photographs of their flowers are available in Vanderplank (2000) and Ulmer & MacDougal (2004).
[Note: this key was revised 29 March 2009, after publication.]
We thank Eva Grotkopp, Fred Hrusa, Rob Klinger, John M. Randall, Bill Sykes, and Peter Williams for many helpful comments.
This is a delightful and impressive book; the result of 13 years of dedicated work. With the objective of providing an up-to date account of all taxa of mosses recorded in the Antarctica, the authors examined the type of every taxon described from the study area as well as many historical collections. The introductory chapters provide information on the biogeographic zones, the climate and geology of the region, a history of bryological studies, as well as a discussion of the terrestrial ecosytems including extreme habitats such as geothermal vents and deep lakes.
The flora is based on traditional taxonomic concepts and therefore is primarily based on morphology. An overall key to the genera is provided, as well as separate keys for the genera for each family and the taxa for each genus. Generous detailed descriptions are given for each taxon with information on morphological characteristics, chromosome numbers, reproduction, habitat, distribution, and similar species.
As the title promises the flora is richly illustrated with detailed line drawings and distribution maps for all species and varieties, as well as forty-two color plates of typical moss communities. The line drawings are delicate and clear. A full page of line drawings are provided for each taxon, including habit sketches, leaves, stem and leaf cross-sections, leaf areolation, gemmae, perichaetial leaves, and in many cases sporophyte characteristics.
This book, along with the previously published Liverwort Flora of Antarctica (Bednarek-Ochyra et al. 2000) and the Lichens of Antarctica and South Georgia: A Guide to their Identification and Ecology (Ovstedal & Lewis Smith 2001), completes a series on the bryophyte and lichen systematics of the Antarctic region; a important addition for an area in which bryophytes and lichens dominate the vegetation.
Related titles:
1a Hypanthium (floral tube)absent or saucer-shaped and inconspicuous; corolla white;
leaves on adult shoots 5- to 9-lobed .................................................................... P. caerulea
1b Hypanthium elongate; corolla pink, magenta, or red; leaves on adult shoots 3-lobed.
2a Sepals longer than the hypanthium; corona filaments in 3 or 4 series, outer two blue 0.2-0.4 cm,
inner white; corolla bright red/scarlet ............................................................. P. manicata
2b Sepals about the same length as the hypanthium or shorter, corona filaments in <4 series.
3a Hypanthium about the same length as sepals; corona filaments in three series, outer 0.6-1.0
cm, of filiform, violet elements, second series 0.1-0.3 cm, membranous, upper 1/2 split into
filiform segments, white, third series 0.1 cm, membranous, white; corolla dark pink ......... P. ‘Coral Sea’
3b Hypanthium longer than the sepals; corona filaments in one series (reduced to a denticulate
ring of teeth in P. tarminiana); corolla pink, magenta, or red.
4a Stipules 3-7 x 1-3 mm, early deciduous; nectary chamber 1.2-2.0 cm wide; hypanthium/
sepal length ratio 1.3-1.6; perianth lobes reflexed, light pink; fruit fusiform;
leaves glabrous to glabrate on upper surface ............................................ P. tarminiana
4b Stipules 6-20 x 12-30 mm, persistent; nectary chamber < 1.4 cm wide; hypanthium/
sepal length ratio > 1.6; perianth lobes campanulate, corolla pink to magenta or
bright red; fruit obovoid to oblong; leaves glabrous to densely pubescent on upper surface.
5a Bracts usually connate for >3/4 length, densely pubescent with conspicuous hairs;
flowers salmon-pink to bright red; hypanthium moderately to densely pubescent;
hypanthium/sepal length ratio 1.4-2.6; leaves glabrous to glabrate on upper
surface; petiole with 4-10 glands ....................................................... P. mixta
5b Bracts connate for 1/3-2/3 length, sparsely to moderately pubescent, with
inconspicuous hairs; flowers pink to magenta; hypanthium glabrous; hypanthium/
sepal length ratio 2.4-3.2; leaves glabrous to glabrate or moderately to dense
pubescent on upper surface; petiole with 6-14 glands ............................... P. tripartita
6a Leaves moderately to densely pubescent on upper surface ................... var. mollissima
6b Leaves glabrous to glabrate on upper surface.
7a Leaves incised ca. 5/6 of their length, lobes narrowly lanceolate ....... var. tripartita
7b Leaves incised ca. 4/5 of their length, lobes lanceolate-eliptic ........ var. azuayensis
Acknowledgements
References
Figure captions for supplementary data at URL: http://bomi.ou.edu/ben/406/passiflora_figures.pdf
BOOK REVIEW: THE ILLUSTRATED MOSS FLORA OF ANTARCTICA
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