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THE BROMELIAD SOCIETY BULLETIN

The Bromeliad Society Bulletin is the official publication of the Bromeliad Society, a non-profit corporation organized in 1950. The Bulletin is issued six times a year. Subscription to the Bulletin is included in the annual membership dues. There are four classes of member-ship: Annual, $5.00; Sustaining, $7.50; Fellowship, $15.00; and Life $100.00. All memberships start with January of the current year. For membership information, write to Mrs. Jeanne Woodbury, 1811 Edgecliffe Drive, Los Angeles, California 90026. Please submit all manuscripts for publication to the editor, 647 South Saltair Avenue, Los Angeles, California 90049

OFFICERS
PresidentDavid Barry, Jr. Editorial SecretaryVictoria Padilla
Vice PresidentFritz Kubisch Membership SecretaryJeanne Woodbury
Treasurer           Jack M. Roth

Board of Directors
Warren Cottingham
Ralph Davis
Nat De Leon
James N. Giridlian
J. G. Milstein
Julian Nally
W. R. Paylen
O. E. Van Hyning
Charles A. Wiley
Wilbur G. Wood
Dr. Russell Seibert

Honorary Trustees
Adda Abendroth, Brazil
W. B. Charley, Australia
Charles Chevalier, Belgium
Mulford B. Foster, U.S.A.
C. H. Lancaster, Costa Rica
Harold Martin, New Zealand
Richard Oeser, Germany
Raulino Reitz, Brasil
Walter Richter, Germany
Dr. L. B. Smith, U.S.A.
Marcel Lecoufle


THE PICTURE ON THE COVER -

This handsome bromeliad is Aechmea ramosa × Ae. weilbachii var. leodiensis. It is one of the very fine Aechmea hybrids that have been made by Mr. Nat DeLeon of Miami, Florida. Mr. DeLeon writes: "The ramosa that I used is what I consider to be the finest of the three forms. This is not the ramosa described in Bob Wilson's book, but rather the type that has brilliant red clasping scape bracts and nearly white berries. This form is late fall to winter flowering and stays in color for a very long-time. While most of the plants of this cross resemble ramosa in foliage color, some have the coloring of the weilbachii. Some of the plants have a tendency to branch more than others also." The editor's plant is one of the most spectacular in her collection. Photo by J. Padilla.


Articles and photographs are earnestly solicited by the editor. Length is no factor. Please mail all copy to the editor, 647 South Saltair Avenue, Los Angeles, California 90049.


NATURAL AND CULTIVATED BROMELIADS IN SOUTHEASTERN BRAZIL

EDWARD L. McWILLIAMS

Author

From January 20 to February 20, 1968, Dr. Lyman B. Smith and I collected and studied bromeliads and visited with bromeliologists in the states of Rio de Janeiro, Sao Paulo, Guanabara and Para. The following is an account of some of our observations and experiences in southeastern Brazil. Figure 1 shows the major area on which we concentrated.

Figure 1. Map of the area collected intensively between Rio de Janeiro and Sao Paulo.

The eastern margin of the Brazilian highland descends abruptly to the sea and forms a steep slope called the Great Escarpment. From Cabo Frio southward the Great Escarpment is unbroken by rivers; but northward from Cabo Frio and Rio de Janeiro a succession of rivers have cut back across the highland. The rain which falls in Sao Paulo flows west and south all the way to Argentina before reaching the sea.

Although there is no true coastal plain like that in the Eastern United States, there is instead a zone of varying width of sand and boulders. The vegetation of this area is called "restinga". There are many miles of white sandy beaches and sand dunes occasionally interrupted by rocky slopes or swamps.

Southeastern Brazil is one of interest to students of the Bromeliaceae for several reasons. Over one quarter of all of the bromeliad species of the world occur in this area. More important than sheer numbers, however, is the fact that in the coastal forest of southeastern Brazil the Bromeliaceae have found an important center of distribution and evolution and many genera and species are endemic to this area. The Bromelioideae, a taxonomically difficult and complex sub-family, is particularly successful here, and important answers to questions of the evolutionary relations within this group may well be found in this area of southeastern Brazil. It is interesting to note that the center of distribution and speciation of many of our best-known ornamental bromeliads is in and around the state of Guanabara. Although the taxonomy of the bromeliads has been studied intensively and extensively by three great monographers, Carl Mez (1891), J. G. Baker (1889), and Lyman B. Smith (1955), surprisingly little is known of their ecology and biosystematics. There are five major papers on their ecology, the oldest of which is Schimper's 1888 paper on the epiphytic vegetation of the Americas. Haberlandt (1914) discussed the physiology and anatomy of many bromeliads, and Picado (1913) made detailed studies of the organisms which live in bromeliad tanks. More recently, the ecology of bromeliads has been of interest for the mosquitoes that breed in their tanks. In 1948 Pittendrigh studied the ecology of the bromeliads of Trinidad in an effort to find ways to control the spread of malaria. The most recent ecological studies of the Bromeliaceae in Brazil have been carried out by Veloso (1952, 1953), upon species native to the states of Santa Catarina and Rio Grande do Sul.

One type of investigation which has yet to be exploited is the determination of whether there are races or ecotypes within the wide-ranging species of bromeliads. From our knowledge of population genetics of temperate plants we would predict that there were. In what ways the genecological relationships of tropical species differ from those of temperate species is still an unanswered question. Studies should be launched to explain the pattern of intra- and infra-specific ecological, physiological and morphological adaptations, and to elucidate the mechanisms by which these adaptations were achieved.

When we arrived in Rio de Janeiro we were met by Dr. Edmundo Pereira and Dr. Armando Huerta. We then visited the Jardim Botanico where Dr. Edmundo has his living collection of bromeliads. Later on the afternoon of our arrival we collected plants in the restinga at Jacarepagua. There we observe huge populations of the showy Neoregelia cruenta and Vriesea neoglutinosa growing in the sterile white sand of the restinga. The Neoregelia cruenta that we observe around Rio was typical of the species as originally described, and they showed very little marmorate spotting. However, 100 kilometers south of Rio all of the Neoregelias we observed were the heavily marmorate plant known as N. marmorata, or intermediates between it and the traditional N. cruenta, the two species intergrading completely. After seeing so many intermediates we feel that there is a real question of whether these two very familiar "species" are actually distinct.

While collecting along the cliffs at the end of Jacarepagua we found Vriesea goniorachis and Dr. Edmundo pointed out that although this species has secund flowers in the living state, when the type was pressed its flowers were spread back and this caused the species to be described as "non-secund". Growing with the V. goniorachis on sheer rock walls was V. reginae in full flower (Figs. 2, 3).

Figure 2. Vriesea reginae growing on steep granite mountain side at Jacarepaqua, Rio de Janeiro.

Collecting in the Parque National de Tijuca we found numerous plants of Quesnelia marmorata with humming birds flying around the flowers. These plants along with Aechmea fasciata were usually growing up in the trees, and if it were not for careful work with Dr. Edmundo's "bromeliotome" we would have had difficulty pulling down these epiphytes (Figs. 4, 5).

We observed many specimens of Billbergia zebrina, all of which were growing in the knot holes left in trees as a result of fallen lower limbs. These plants were often difficult to collect because their basal parts were embedded one to three inches inside the tree. B. zebrina is striking for its habit of growing out at an angle of about 45° from the tree. It is largely limited to this "knothole" niche, at least around Tijuca. Observations on the germination and establishment of B. zebrina in nature would be interesting. Possibly birds which eat the fruits of the plants use these tree holes as nesting places and the plant becomes dispersed and established in this fashion.

While collecting along the road up the Corcovado (a 2,000-ft. mountain distinguished by its 100-ft. statue of Christ at the crest) we discovered Vriesea saundersii growing with Vellozia candida (Velloziaceae) on steep granite cliffs.

Figure 3. Vriesea regime, V. goniorachis, and Tillandsia araujei growing on granite mountain side near Rio.

After working around Rio we devoted several days to the area of Teresopolis. In the Park of the Organ Mountains, or Parque Serra dos Orgaos, we found a large hybrid population of Vrieseas growing along a roaring stream. On one side the trees were loaded with what appeared to be Vriesea carinata × ensiformis hybrids, and on the other side were V. carinata × scalaris. In the same area we found two other species, V. guttata and V. platynema, the latter being very common throughout much of coastal southeast Brazil.

A few kilometers south of Teresopolis we passed a small roadside flower stand which contained another Vriesea hybrid (Fig. 6). This appeared to be a V. psittacina hybrid. The boy who was selling the plants said that they were harvested locally, but the exact location seemed to be a trade secret.

In Teresopolis we visited Mrs. Adda Abendroth and spent an afternoon in her bromeliad garden, where we saw numerous Vriesea species, including some she has recently discussed in the Bromeliad Society Bulletin (Abendroth, 1967). Among the many interesting species that she cultivated were several races of Tillandsia stricta which she had carefully studied for several years. One of the races consistently produces flowers in August each year, while another of the races flowers only in February. This is very interesting from an ecological point of view as genecological studies of temperate plants have long demonstrated the existence of races or ecotypes of plants which flower at certain times of the year (Turesson, 1925).

The question of how and why certain species are common and wide-ranging and others are rare and have limited ranges remains to be answered for most tropical plants. Bromeliads and other epiphytes are particularly appropriate for ecological studies. Their distribution is cleanly correlated with ecological factors such as microclimates and substrates. Richards (1964) has pointed out that the extraordinary structural specializations of such epiphytes as bromeliads—more perhaps than those of any other group of plants—truly deserve to be called adaptations.

The following week while on our way to visit the garden of Mrs. Ruby Braga in Petropolis we passed several trees laden with Quesnelia edmundoi and species of the epiphytic cactus Rhipsalis. We often observed epiphytic cacti growing in clumps of bromeliads. The bromeliads apparently act as colonizers, and after they are established other kinds of plants germinate in the soil that accumulates around the bromeliad roots.

In Mrs. Braga's garden we saw numerous unusual bromeliads, including several which Dr. Smith was not able to name. One of these appeared to be a new species of Wittrockia, but unfortunately the plant was not in flower. Mr. and Mrs. Braga later took us to a rainforest a few miles from Petropolis, where we saw about twenty bromeliad species growing in a single acre of ground near a stream. Among the plants were large clumps of Neoregelia tristis on moist rocks and Nidularium fulgens with its blue petals and nest-like inflorescence. In the trees on the hill high above the stream were growing several Vriesea species, including V. procera and V. phillippocoburgii (Fig. 7).

Our next campaign was in the area of Angra da Reis. Coming down the mountain overlooking the Baia da I. Granda we traveled over an old cobblestone "paralelipipido" highway. About 15 km. east of Angra da Reis we collected in a marsh a few kilometers from the coast. Smith (1962) has pointed out that for thousands of miles one can pick a stretch of Brazilian coast at random and know that he can find Rhizophora mangle, Hibiscus tiliaceus, and other tropical maritime species. Numerous bromeliads were present in this marsh, but most of them were epiphytic on shrubs which were just out of reach of our "bromeliotome". Several of the bigger ones that "got away" were Quesnelia quesneliana, Q. arvensis and a Tillandsia species. Later we waded out into the marsh, attempting to stay on the tussocks of grass to keep from sinking into the mud. One specimen of Vriesea procera that we inspected had a green snake coiled up in the tank which departed when we attempted to collect the plant.

In general the best plant collecting is along the poorest roads, but the old road that leads up the mountain from Angra da Reis proved to be an exception. About ten kilometers along the road up the Great Escarpment we stopped to collect a large Aechmea that was growing on a rock outcrop above the highway. The first swing of the bromeliotome missed the Aechmea but came down with a smaller bromeliad which we had not seen. This plant had long rhizomes and yellow-orange leaf apices and flowers. The other plants of the clone were out of reach and we had to climb the cliff to get at the plants. "Unfortunately", (or "infelizment", a very popular word in Brazil), the rocks were quite slick and I slid back to the base of the cliff several times before getting to the top. There we found several common bromeliads growing with the unusual plant, including Neoregelia tristis; Pitcairnia flamea, and Nidularium innocentii.

After much dissecting of flowers and pacing to and fro Dr. Smith decided that our plant was a new species of Wittrockia, but it is quite different in habit and growth form from most Wittrockias. This species may prove to be of interest to horticulturists as it is rather ornamental.

On the night of February 1 we stayed on the bay of Grand Island. The view from the shore was very striking, with many small islands dotting the blue Atlantic as far as one could see. There are hundreds of these islands along the coast of Brazil. These are continental islands for the most part, located on the continental shelf. The sea is very shallow along this stretch of coast and at some time in the past no doubt these elevated land areas were connected to the mainland. The flora of these islands has not been studied extensively, and it is likely that some of the more isolated islands have new bromeliad species on them. Islands have always held a particular fascination for the student of plant geography and evolution. Indeed, many of Charles Darwin's ideas were based on observation of the fauna and flora of islands off the coast of South America.

The following morning we attempted to call the Instituto de Botanica in Sao Paulo. The operator greeted us with the now familiar "infelizmente" it will take several hours to get the call through. Since several hours can easily become a day in Brazil we decided not to wait.

In Sao Paulo we stayed at the Instituto de Botanica through the courtesy of the Director, Dr. Alcides R. Teixeira. The gardens were excellently landscaped and in good condition. In the natural areas section of the garden we saw several monkeys and parrots, along with numerous bromeliads. We spent the following day visiting Dr. and Mrs. Eiten, who are both botanists at the Botanical Institute. Among the interesting plant collections there was a good number of bromeliads which Dr. Eiten has collected in his ecological and taxonomic studies of the vegetation of Sao Paulo State.

On Saturday afternoon we visited the garden of Mrs. Peggy Pollard. She had a variety of bromeliads under cultivation and one of her most interesting plantings was a bed of Neoregelia compacta (Fig. 8). We were later able to collect this plant along the seashore at Parati and in the bromeliad garden of Dr. Burle Marx.

From Sao Paulo we traveled southeast down the Great Escarpment, passing through very scenic country. The eastern margin of the Brazilian highland descends abruptly to the sea. In some places the descent is over a "series of steps, but at the back of Santos there are some places where the descent from 850 meters to sea level is accomplished over one steep slope. This sharply marked off eastern edge of the highland is called the Great Escarpment. Between Santos and Sao Paulo it was given the name Serra do Mar, although it is not actually a mountain (serra). Geologists consider the Great Escarpment to be a monoclinal flexure rather than a fault scarp. The vegetation of the area was interesting and at times we saw whole mountain sides covered with several species of Tibouchina (Melastomataceae) in full flower. Dr. Smith collected seeds and specimens of some of these for Dr. John Wurdack of the Smithsonian Institution, who is an authority on the family.

A few kilometers north of Santos on the Ilha Santo Amaro Guarija Tillandsia stricta and Vriesea rodigasiana were common in the low trees and shrubs of the coastal restinga. We took the balsa across the bay to the small town of Bertioga and then drove north along the sandy seashore for about 30 kilometers; this was one of the smoothest roads we traveled while in Brazil. Soon after we passed through Bertioga we began to see several different marmorate bromeliad species. Quesnelia marmorata, Neoregelia marmorata and Vriesea corcovadensis were common, and often all three species were growing together. We also saw numerous plants along the beaches which appeared to be intermediate between Neoregelia cruenta and N. marmorata. We brought back living plants of these and hopefully chromosome and progeny studies will provide some explanation for the observed variation. Possibly introgressive hybridization is involved. As we moved up the coast we would occasionally have to cross rocky headlands, and the vegetation would abruptly change. Quesnelia arvensis, Aechmea pectinata, and A. nudicaulis often grew on boulders which were splashed by the surf. The populations should prove to be interesting in genecological studies of bromeliads as they must have a high tolerance to the salt water that accumulates in their leafcups.

About half way between the island of S. Amaro and the town of S. Sebastiao the bromeliad flora of the restinga seemed to change, and we began to find Catopsis berteroniana, Vriesea incurvata and V. fenestralis in the low growing trees. One advantage of collecting epiphytes along the coast was that none of the bromeliads were out of reach of Dr. Edmundo's five meter bromeliotome. In the sandy soil along the beach Ananas fritzmuelleri was common. This is about as far north as this species goes. Another maritime terrestrial species which we saw along hundreds of kilometers of coast was Bromelia antiacantha. This is a wide-ranging species and it grew in more open situations and not in partial shade where Ananas fritzmuelleri was found. We collected seeds of several bromeliads, and they all had seeds that were completely dormant. One wonders if this seed dormancy is an adaptation to the severe environment in which they grew. Following a "wild" ride over the headlands to Sao Sebastiao we arrived in the city about midnight.

The following day, February 6th, we backtracked southwest 20 to 30 kilometers in order to see some of the country we had driven through the night before. Our first stop that morning was at a large cluster of boulders near a small bay. Here, to our surprise, we found a number of crabs living in the tanks of the Neoregelia cruenta which grew on the granite rocks. The crabs are presently being identified at the Smithsonian Institution. A few kilometers west of the bay we found large populations of Vriesea ensiformis in the trees which grew along the banks of a stream. This species was quite common immediately over the stream, but 10 to 20 meters from the rushing water the plant was not present. It was quite easy to see the distribution of almost every Vriesea as the woods were fairly open and the bright red inflorescences stood out like torches. Apparently the high humidity near the stream in this area is critical for the survival of V. ensiformis. In the rock crevices along the stream we found Aechmea coelestis and Nidularium burchellii.

Left—Figure 4
Dr. Smith contemplating pressing the afternoon's haul.

Below—Figure 6
Vriesea psittacina hybrid in road-side flower stand.

Author
Figure 5. Dr. Edmundo coaxing a Vriesea out of a tree at the Parque Serra dos Orgaos.
Author
Figure 7. Mrs. Ruby Braga and a large specimen of Vriesea phillippocoburgii collected near Petropolis.

Going north, again, toward Caraguatatuba the vegetation changed to coastal swamp, which provided rich bromeliad hunting. We visited one interesting swamp about 20 kilometers northeast of Caraguatatuba where we collected a form of Nidularium seidelli which has red bracts. In one area I waded out into the water to collect what I thought to be an unusual Billbergia. After climbing a small tree growing in the knee-deep water I discovered that my plant was only the common Quesnelia marmorata. The compound pendent inflorescence with purple petals closely resembles a Billbergia from a distance. The next turn in the road presented several Quesnelias that could have been collected quite easily. We spent the night in Caraguatatuba in a hotel near the coast. There have been several desastroso landslides in this town and we were careful to choose a hotel well away from the cliffs.

The following day we stopped at the mouth of the scenic Rio Escuro on the road to the coastal town on Ubatuba. Here again we found numerous crabs living in the leaf bases of several bromeliads. The trees and rocks along this bay were literally covered with bromeliads. On the shaded rocks we found Neoregelia ampulacea, Aechmea pectinata and Nidularium procerum. In the trees were many plants of Vriesea guttata in full bloom and on the rocks in full sun and near the breaking surf grew Catopsis berteroniana and several forms of Vriesea procera.

We arrived in the quite and sultry little town of Ubatuba at about four in the afternoon. As usual, against better advice, I optimistically thought we could make it to the next town before dark. As we slowly ascended the mountain road to Sao Luis do Paraitinga we passed through an interesting succession of bromeliads. On the rocks along the coast were the ever present Neoregelia cruenta, Aechmea nudicaulis, Vriesea procera and in the restinga V. platynema was present but not common. At about 300 meters we stopped seeing the first three species but V. platynema became quite common in the trees. Several Billbergia species and Canistrum lindenii were also evident at this altitude. At 600 meters Vriesea platynema was no longer present but Nidularium purpureum was common on the moist, shady sides of boulders. At the crest of the Serra, which is approximately 1000 meters, most of the coastal species were completely absent. Although the clouds were very thick and visibility was limited to 30 meters, there was enough light to see hundreds of plants of Vriesea altodaserrae; this species was clearly the dominant bromeliad on the mountain top. In the rich leafmold and moss beneath the Vriesea-laden trees we found masses of Nidularium innocentii, but the most interesting find was a large Vriesea with green triangular floral bracts which appears to be new to science.

The afternoon was well advanced and we had many kilometers to travel before night, so we did not have time to collect this mountain crest carefully, but further exploration of the area should produce some interesting bromeliads. The road had long since become a rocky trail and we made slow progress getting down to Sao Luis. As we bumped down the mountain in the dusk we could see the silhouette of groves of Aracauria angustifolia against the skyline. These were some of the largest Parana pines that we saw. Unfortunately there are relatively few stands of this species left in this part of Brazil and we did not see any signs of reforestation programs. The destruction of tremendous areas of natural forest in southeastern Brazil has resulted in extensive erosion and numerous landslides. West of Caraguatatuba we saw all the vegetation on a mountain go up in smoke in one night. The rainforest burned slowly but at very high temperatures and it took about eight hours for the fire to progress from the base of the mountain to the crest. In another area near S. Sabastiao we saw fires, overgazing and landslides all on one large serra.

The next day was bright and sunny and we had hopes of bigger and better bromeliads, but we saw little forest worth collecting in as the agricultura had swept the country clean. Since we had found new species both to the north and south of this area, one can be pretty sure that some of the bromeliad species of this country had been destroyed before they were ever collected. Irwin (1967) working in central Planalto part of Brazil noted that botanists are hard put to complete preliminary studies of the tropical flora before civilization eliminates rare plant species in the relentless conquest of nature. We later observed similar destruction of the natural vegetation in northeastern Brazil near the Amazon. That afternoon we returned to Rio to process our plants for shipping.

We spent the next few days recuperating from our trip and visiting with botanists in and around Rio. At the Museu National we spent the day with Dr. Segadas-Vianna and were impressed by his detailed Flora Ecologica de Restinga project. Dr. Segadas coordinates the work of a number of scientists in their study of the coastal vegetation. Included in the ecological laboratory of the National Museum are taxonomists, ecologists, zoologists, chemists and geologists, each studying the restinga from a slightly different point of view. Of the 45 bromeliad species that grow in the coastal vegetation relatively little is known about their ecology and there is muito ecologica work that needs to be done on the family.

By now we had collected along most of the coast between Rio de Janeiro and Santos, the port of Sao Paulo, but there was one area in the vicinity of Parati that we had not collected. On Monday February 12th we left Rio and traveled south to the town of Cunha from the isolated coastal town of Parati. A few kilometers east of Parati we found a campo or "prairie" type of vegetation which reminded me of Iowa. Most of the same genera which one sees in a North American prairie were represented and the most important family was the Graminiae. There were also sedges and Compositae, including one large Senecio spp. and a spectacular two meter Lobelia.

It was interesting to observe the change in the bromeliad flora from the dry western side of the range to the moist eastern slope. The flora is always richer on the seaward side due to the increased precipitation.

Figure 8. Dr. Smith and Margaret Mee examining Neoregelia compacta in the garden of Mrs. Peggy Pollard in Sao Paulo.

Figure 9. Dr. Marx, Dr. Smith, and Dr. Edmundo in the bromeliad collection of Dr. Marx.

We spent most of the following day exploring the countryside around Parati and we found several interesting plants. Approximately eight kilometers north of Parati we found Neoregelia compacta growing in partial shade on a granite boulder. In the same area we photographed a large tree that was literally covered with Vriesea procera and Neoregelia cruenta, both in full flower. These two species along with Aechmea distichantha and Tillandsia usneoides were common along the coast. However, the festoons of Spanish moss never approached the size they attain in the southern United States. At an altitude of 300 meters above Parati we began to see Vriesea scalaris dotting the older tree limbs and near the crest of the serra V. guttata and Aechmea pineliana were common.

The following day we moved north and collected along the coast and in the valleys around the old town of Mangaretiba. Unfortunately progress and civilization had beaten us to the vegetation again; this time we were 400 years too late. We traveled along a crumbling cobblestone road through valleys that had been producing bananas on and off for 400 years. Occasionally we would see coffee trees growing along the side of the road, a relic of a once thriving coffee plantation. There is supposed to be a mountain "road" that connects Mangaretiba and Angra dos [check!] Reis, but it is strictly a dry weather road and it was beginning to rain when we finally found this road. A collecting trip along this trail might turn up some interesting bromeliads.

One of the bromeliad gardens which we had hoped to see was that of Dr. Burle Marx, and the day before we left for Belem we were fortunate enough to visit the estate. Dr. Marx is an internationally known landscape architect and an excellent horticulturist as well. He has a large bromeliad collection (Fig. 9) with numerous naturalistic settings. Among the many interesting species were two Hohenbergias which Dr. Edmundo and Dr. Smith had never seen before; this is a pretty good sign that no-one else has ever seen them either. Specimens of these plants are being studied by Dr. Edmundo. Hidden among the hundreds of Heliconias in the garden was a two meter specimen of Aechmea blanchetiana in full flower. This species is very similar in appearance to a Vriesea. Dr. Marx and Margaret Mee have found many interesting bromeliads in northeastern Brazil, including Encholirium spectabile, Orthophytum saxicola and several Cryptanthus species. These and other saxicoles were growing in sandy soil on a dry rock outcropping in the garden. Dr. Marx gave us several Aechmeas, Billbergias and Pitcairnias for the University of Michigan Botanical Gardens' bromeliad collection and we promised to send him plants of our new species.

The following day, February 17th, we took a flight to Belem, Brazil, for several days of collecting near the Amazon. The bromeliads which we collected during this field expedition in southeastern Brazil include over 125 species, many of which we were able to get back to Ann Arbor alive. Of this number two new species and several others were either new species or varieties. A number of populations of some of the wide-ranging bromeliad species were collected in various geographical areas, and these should prove interesting in future ecological studies of these plants. Field observations of several hybrid swarms will be followed up with cytological and anatomical studies which, hopefully, will identify the parental species of the hybrids. We also collected many propagules and herbarium specimens of other plant families, including the Melastomataceae, Gesneriaceae, Rubiaceae, Myrtaceae and Malpigiaceae.

—Horticultural Botanist, University of Michigan Botanical Gardens, Ann Arbor, Michigan.

BIBLIOGRAPHY

Abendroth, Adda. 1967. "Vrieseas around Terespolis, Brazil", Bromeliad Soc. Bulletin. XVII: 54-63.

Baker, J. G. 1889. Handbook of the Bromeliaceae, London.

Harberblandt, G. F. J. 1914. Physiological plant anatomy. MacMillan, London.

Irwin, Howard S. 1967. "A Year in the Planalto of Brazil". The Garden Journal. 17: 46-59.

Mez, C. Bromeliaceae. 1935. In Engler, Das Pflanzenreich, 103 (LV): 32-100.

Picado, C. 1913. "Les Bromeliacees epiphytes, considerees comme milieu biologique. Bull. Scientif. France et Belgique. 47: 215-360.

Pittendrigh, Collin S. 1948. "The Bromeliad-anopheles-malaria Complex in Trinidad. I-The Bromeliad flora. Evolution 2: 58-89.

Richards, P. W. 1952. The Tropical Rainforest. Cambridge University Press, Cambridge.

Smith, L. B. 1955. The Bromeliaceae of Brazil. Smithsonian Miscellaneous Collections. 126: 1-290.

Smith, L. B. 1962. "Origins of the Flora of Southern Brazil". Contributions from the U.S. National Herbarium. 35: 215-249.

Turseson, G. 1925. "The Plant Species in Relation to Habitat and Climate". Hereditas. 6: 147-236.

Veloso, H. P. 1952. "O problem ecologico vegetacao-Bromeliaceae-arofelinos." Anais Botanicos. 4: 187-228.

Veloso, H. P. 1953. "O problem ecologico vegetacao- Bromeliaceae-anofelinos. II. Avaliacao quantitative dos criadores". Anais Botanicos. 5: 5-36.


BROMELIADS-HOUSEPLANTS FOR TODAY AND TOMORROW

WALTER RICHTER

(Translated from the German by Mrs. Adda Abendroth)

MULTIPLICATION BY SEED

First-rate seed is, of course, necessary for good results because the development from grain to full-fledged plant is long. Therefore, those who are anxious to avoid the risk of having worked in vain should use only fully guaranteed seed. The viability of seeds is short, six months from the time of ripening being the limit. The date of collection does not necessarily coincide with ripeness. Some species retain ripe seeds in the capsules or in their berries for weeks or months without getting dispersed. It is useless to plant seed that is over six months old. Occasional assertions heard to the contrary do not hold. There are three ways to secure first-class seed. The surest, and also the most time consuming, is to grow it yourself. This method calls for a sufficient number of mother plants and pays off when growing on a large scale only.

Purchase seed only from reliable dealers, those who can furnish viable seed that has recently been collected. If a large quantity of seed is to be bought, it is well to order it beforehand so that it can be delivered immediately upon being collected. On the other hand, it is reasonable to be lenient if immediate delivery cannot be made—bromel seed in perfect condition is not an item to be always in stock. The third way of producing seed is importing it from its country of origin. This may seem a comparatively safe way, but in reality it is not. Although air transportation is no problem nowadays, the distance from the habitat of the plant to Europe is long. The dealer abroad depends upon the collectors who do not always understand what is at stake.

As I have said, the final phase of development of capsule fruits is often very rapid; within a few hours the seeds are gone. Harvesting from wild plants is thus more often than not a matter of good luck; in addition the capsules on one spike don't all ripen at once. Whether or not post maturation on cut-off spikes guarantees first-quality seed is yet to be proved. My misgivings concerning imported seed, however, should not be taken as total condemnation. I have derived much pleasure from high-percentage germination in imported seed but total losses have brought deep disappointment. In any case, imported seed should be planted immediately upon receipt, not a day should be wasted. For some of the species that take a long time to flower, such as Vriesea hieroglyphica, V. fenestralis, and others, we depend almost entirely on imported seed. The importation of seed is also the best way to introduce newly discovered species. It is hoped that as the demand increases, growers in South and Central America will raise their own bromels for seed and that the quality of the seed produced will improve.

The question as to the best time to plant seed depends less on our own personal notions and preferences than on the availability of fresh seed in perfect condition. It is advisable to plant all seed that is purchased immediately. I plant seed the year round independent of season. If the surroundings are warm enough, there is no difference whether the seed is planted in summer or winter. Floss-appendaged seeds do not require much light during the first weeks after germination, so that absence of brightness in our northern winter months hardly affects the seedling.

Planting techniques vary with the different species. Grains coming from berries, such as Aechmea, Billbergia, Cryptanthus, Neoregelia, Nidularium, pose no problem. The granules should be strewn in moderate density on a mixture of well-seasoned leafmould and finely ground peat and sand. The compost must be loose and spongy, never compacted. Proper aeration is most important. Other mixes for instance, heather-soil or pure, not too-acid peat mixed with sand, are good for germination, provided their ph value is around 4. For containers, perfectly clean, common pots can be used, round or square earthen bowls, depending on the amount of seed to be planted.

Seeds usually retain residues of the sugary pulp that covers them in the berry. Its presence favors the development of mildew, which in turn destroys viability; therefore, seeds should be disinfected before planting. I see to it that disinfection is carried out when the seed is being collected. Squeeze the berries between two fingers and let the content drop into shallow glass bowls containing a little water. Keep them in a warm place for 2 to 4 days. Fermentation will then free the grains from the membrane that enfolds them and from any remnants of pulp. Rinse the seeds thoroughly, using a fine sieve, and put them in a disinfecting solution, such as Chinosol 1:1000 or Sulfachin 1:1000. Make sure that all grains get immersed. Then they may be spread to dry. As seeds offered for sale are not always disinfected, it is well to do the disinfecting yourself, either before or after planting the seed. I usually get all seed containers sprayed generously with a disinfectant as soon as the seed has been distributed, enough to get the mixture soaked. The grains must be left uncovered; nearly all bromels need light to germinate.

In order to keep the seeds from drying out during the first few days of germination, cover the containers with a piece of glass, allowing for some ventilation. Species with berry-fruits and distinct kernels germinate rapidly. Germination depends on temperature; it should not be too high, 20° C average being sufficient. The seed containers should be placed in a well-shaded house as near the glass as possible. Constant and even, not too much, humidity is essential. Excess moisture endangers success, especially at the point when the cotyledon is out and the seedlings are too close together to get rid of excess moisture before night sets in. After a fortnight, the glass is usually no longer needed. In order to harden the seedlings see that additional ventilation is provided as the days grow warmer. Seedlings treated in this manner may be transplanted after 4 to 6 weeks.

The planting of seeds with a hair-crown is considerably more difficult. These would include Guzmanias, Vrieseas, and Tillandsias. Their seeds are collected dry and need no prior disinfecting. The same type of dishes may be used as described above.

Ordinary square or rectangular day containers are disinfected by steam. In the bottom place a layer of coarse, steamed peat about 2 cm thick; press it down. Next comes a layer of chopped osmunda. Compact it, pressing lightly, leaving about 2 cm free below the rim. Now spread the seed. Use a pair of pincers to distribute it evenly over the whole surface; don't let the hairs make a clump, as algae might form. This method approximates the procedure of nature in the plant's homeland. The layer of osmunda provides good ventilation; the highly moisture-absorbent layer of peat acts as a source of rising humidity, thus constructing an excellent micro-climate in a very small area. The planted seed is not to be covered. The next step is a shower of disinfectant as described for the first group. Apply glass plates in the same manner. The containers go to the warmest house—22-25° C or even warmer. The seeds could germinate even at 20° C but could take considerably longer.

Germination occurs within 15 to 25 days, depending on season, site, and warmth. It is much slower than in the former group. During the first period spraying should be done very carefully. The moisture that rises from the peat layer provides even air humidity, further aided by the precipitation on the glass plate. I must stress that the top layer with the seeds is not to be damp all the time; a frequent alternation between moist and slightly dry is the correct condition for germinating bromeliad seeds. Even occasional drying will not harm seed in this phase of germination as long as it is only a passing stage and not a permanent condition. Very early in life the tiny seedlings exhibit the hardiness appropriate to epiphytes. Development is very slow, but experiments have shown that bromel seedlings of this group resent contact with fertilizers. In winter I recommend artificial light. The seedlings should be left in their original containers as long as possible as long as the formation of algae or moss does not disturb them. Too early transplanting is not recommended; normally I wait until the plants are 4-5 months old.

For especially valuable seed I recommend the following method: Line the bottom of shallow Petri-dishes with a layer of plastic sponge about 1 cm thick, spread the seed on it and then spray with disinfectant. Discard what fluid may collect in the bottom of the dish. As the dishes are covered, evaporation is avoided; all that is necessary is to give the contents an occasional examination to see if all is well. It is a time-saving process. As the seedlings develop, they need more air.

Let them have it by putting the lid on in a slanting position, leaving an open segment. As the seedlings grow, enlarge the gap, and when the young leaves reach the lid, leave it off altogether.

Transplanting is practically alike in both groups, differing only in timing. For both types of seedlings, which are comparatively small in size, the material used is the same: a finely strained mixture of leafmold and peat and sand—about 2:2:1, or else a sandy heather soil. The scrupulously clean clay containers can be lined with a coarser material of the above combination, putting the strained layer on top. Seedlings of Group I, that is Aechmea, Billbergia, Neoregelia, and Nidularium, that grow faster should be spaced a little farther apart than those of Group II, Guzmania, Vriesea, Tillandsia, which take longer to grow. In the first transplant, it is well to keep the plantlets close together, the seedlings developing better this way.

Transplanting should be carried out with the utmost care, especially in Group II. The little plantings often retain flyhairs attached to them; clumsy manipulation in removing the seedlings from the container can easily damage the young leaves. Roots are extremely few in Group II, while Group I possesses a nice little clump. Each group retains its type of roots for life. After replanting, the surface of the potting mix should not be too compact, for that would favor the formation of algae and mildew. The next step is to proceed under identical conditions as those described for the first period—same temperatures and sufficient shading. The containers should be moderately moist, not soaking. Spray at least once or twice a day to insure humidity. Only when the plantlets speed up their growth rhythm should warmth be reduced.

About 8 or 10 weeks later Group I will need a second transplant into a coarser mixture, provided the plants keep growing happily. Group II takes longer to attain this second state, so that transplanting will come about 3 to 4 months later. Development of plantlets in a given lot is often uneven—a natural characteristic. It is advisable to assemble the different sizes from the first. Bromeliads possess the highly convenient quality of being less tied down to a schedule than other plants. Some of the steps may be postponed from one to three months without damage to the plants, provided, of course, that general conditions remain sound. Young seedlings should never be put into dark corners or exposed to low temperatures.

The principle of proximity in transplants should be followed in successive steps. Favorable surroundings help the plants develop better. By the end of the first year, Group I should be weaned to temperatures from 15 to 20° C and more light. The seedlings will then have reached the stage when it must be decided whether they shall go into pots or into the soil. Group II reaches this stage only within a year and a half. Until then the young plants may be kept in their containers or in boxes on the upper shelves of the greenhouse, or suspended, since the only care they need is copious spraying and an occasional watering. Plants cultivated below them will not be affected, and this is another plus in raising bromels.


A GUZMANIA RESEMBLING A NEOREGELIA

RICHARD OESER, M. D.

In an article describing the genus Guzmania, I came across the name Guzmania sanguinea. No information concerning the plant was given, but the ominous sound of the name, "the bloody one", stuck in my mind and I tried to find out more about this particular bromeliad. To my dismay a search in the literature at my disposal was of no avail. Little did I suspect that I had a specimen of the mysterious subject growing right under my eyes. It was a plant I had raised from babyhood, received in a shipment of unidentified plantlets a friend sent me from Costa Rica. The plant had looked different from the rest since early youth with its stiff dark leaves embellished with reddish striata.

At the end of a rest period in the spring of 1964 I noticed certain changes in my plant. They were a sign that a bloom might appear in the not too far-off future. The rosette spread and flattened, the center broadened making room for more water, and the innermost leaves turned red, bloody-red. It was not the luminous flush of a Neoregelia carolinae or its hybrids when a translucent glow seems to come from within. No, in this bromeliad the red looked more like splashes of paint dabbed on with a brush here and there in irregular patches. Instead of the flower spike I expected to see rising from the plant's heart, the pond soon contained an assembly of long individual buds poking well out of the water and opening into pure yellow flowers. The flowers lasted longer than those of a Neoregelia, which seldom live more than a few hours or a day at the most. Occasionally I counted up to five simultaneous blooms in my cup, the oldest, of course, on the verge of decay. All the same, I managed to pollinate the earlier with the later flowers.

A casual observer seeing my plant among the Neoregelias might well take it to be a member of that genus. At first sight the two look much alike. A closer examination, though, will soon reveal individual differences that place the two plants definitely apart. Wholly unarmed leaves exclude Neoregelia from very beginning. Blood-red stains on the blades and the yellow flowers establish it to be Guzmania sanguinea beyond a doubt. Fruiting in both differs greatly.

Most of us know how seeds develop in Neoregelias: the berries grow and mature in the depth of the pond in the heart and when they are ripe a bird called "tie" as observed by Adda Abendroth pulls them up, squashes and chews them and swallows the pulp, cleaning its bill on a perch after the meal. It is also assumed that seeds having passed through the bird's intestines germinate on branches where they happen to be discarded.

Like all other members of the sub-family Tillandsioideae, Guzmania has capsule fruits that dry as they mature, spring open when ripe, and release parachute-borne seeds to the air. Be it that my efforts at pollinization were successful and that Guzmania sanguinea is self-fertile, it soon seemed as if all the flowers in my plant had set fruits. I could feel them with my fingers in the pond where they continued submerged. How they would some day embark on their flight in the air puzzled me greatly at the time.

As always when Nature solves a problem, the process is ingenious and very simple. Just that also happens in Guzmania sanguinea. One important step toward a satisfactory solution is that the capsules grow to considerable length (4-5 cm). They were soon visible above water level. A second favorable condition is that the new shoot sprouts and develops close to the maturing infructescence, in the leaf-axil right next to it, much like in Vriesea splendens. The growing pup forces the fruit-assembly to the side, the cup tilts slightly, the water runs out. The capsules continue their life in a dry environment. It takes a whole year for the seeds to ripen, for the large pods to split and cast their winged charge to the air. By that time the pup is almost ready to flower. Its leaves spread like a roof above the ripening fruit, protecting the capsules from harmful rain.

By a coincidence I later received Guzmania saguinea also from Venezuela and Ecuador. Some types have very ornamental red striata. Thanks to their fine coloring and conspicuous design on the leaf blades even young plants are little gems that make no extra demands on the grower. They may be grown on wood or in a pot with an airy compost mixture. Cultivation from seed on tree fern slabs hung up horizontally is easy. Seedlings survive dry periods well: perhaps they even need these periods to develop properly. Grown outdoors coloring will probably be more intensive than what our picture shows.

—Germany. (Translated by A. Abendroth)


W. Richter

One of the most charming of all Guzmanias is the European hybrid known as Guzmania × 'magnifica.' This cross of Guzmania lingulata var. minor × Guzmania lingulata var. cardinalis was made by the noted German hybridist Walter Richter in 1937, flowering for the first time in 1939. Ever since its introduction into the trade it has been a popular and sought-after item.

The brilliant star-like bract makes its appearance most appropriately during the holiday season and remains in glowing color well into summer. In a well-developed plant the bract will reach a width of six inches. The dainty appearance of the foliage is due no doubt to the influence of the pollen parent, for the narrow thin leaves of the minor strain are at once evident.

This Guzmania has all the characteristics that a grower looks for in a bromeliad: It is a fast grower and a dependable bloomer; it appears to be pest free; it is very generous with offshoots; and at all times makes a handsome appearance. Unfortunately for some, it needs the warm humid atmosphere of the greenhouse.

—V.P.


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