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The Aroids of the West Sarawak Limestone
The limestone (batu kapur in the local Bahasa Melayu) areas of Sarawak are among the most rewarding for searching for aroids, not only in terms of species richness but also species diversity. A particularly striking aspect of the limestone flora is that spatially close areas of limestone, often only a few kilometres distant, often have strikingly different species composition. Over the past few years we have been investigating three areas of limestone in western Sarawak. One area is the Bau series limestones in Kuching Division. Another is the Padawan series that run through the southern parts of Kuching and Samarahan divisions towards the border with Kalimantan (Map 1). These are in the main areas of often very large exposed karst fossliferous limestones dated to the Jurassic and Cretaceous periods. We have also been studying the Ranchan limestones on the outskirts of Serian. These are very old, in parts Permian, argillaceous limestones and mark the eastern-most block of the western Sarawak limestones. All these limestone areas have high levels of local endemism, notably in the genera Amorphophallus, Alocasia, Schismatoglottis, and Homalomena. Additionally, there are suites of species that while not locally endemic in western Sarawak are restricted to limestones wherever they occur in Borneo.
Amorphophallus is one of the most striking genera of the Sarawak limestones. The area around Bau hosts A. brachyphyllus Hett. (Figure 1), which is replaced by the closely related A. eburneus Bogner (Figure 2) on the Padawan series, and by the recently described A. ranchanensis Ipor, Tawan, A.Simon, Meekiong & Fuad at Ranchan (Figure 3). The last named is particularly interesting in that its relationships appear to lie with the east Bornean, A. hottae Bogner & Hett., rather than with any of the currently known species in western Borneo. However, there exists the distinct possibility that as further work is undertaken in Sarawak yet more species will be discovered that fill the geographical gap between these Western and Eastern taxa. Amorphophallus hewittii Alderw. (Figures 4) occurs on all the western Sarawak limestones and, unusually for a limestone-occurring Amorphophallus, also occurs on other geologies, including sandstones (as on the Matang Massif and across the Sarawak River at Gunung Muan) and granite (e.g., at Gunung Gading). One possibility is that the limestone elements represent a species distinct from the plants on other geologies.
Alocasia is another feature of the Sarawak limestones. Alocasia ridleyi A.Hay (Figures 5 & 6) occurs on all three of the western Sarawak limestone formations, although is commonest on the Bau series. Alocasia ridleyi is one of a number of similar species restricted to limestone in Borneo. In the same group as A. ridleyi is A. puteri A.Hay and A. pangeran A.Hay, both restricted to Sabah while a related species, A. scabriuscula N.E.Br. (Figures 7 & 8) occurs throughout much of Borneo, often in rather swampy places. By comparison, A. reversa N.E.Br. (Figures 9 & 10) is restricted to the Padawan series, and is replaced by an as yet undescribed species on the Bau series.
Both Schismatoglottis and Homalomena are abundant on the Sarawak limestones, although the patterns of species distribution are far from resolved. A striking feature of the Bau and Padawan limestones is a species of Schismatoglottis, S. nervosa Ridl., with ribbed petioles and tissues that are strongly aromatic of terpenes (Figures 11, 12 & 13). At Bau, but absent from Padawan and Ranchan, S. bauensis A.Hay & C.Lee forms large colonies on vertical limestone cliffs (Figure 14) and is readily identified by the long free ligular portion to the petiolar sheath (Figure 15) and bright pink innovations (Figure 16). Schismatoglottis calyptrata (Roxb.) Zoll. & Moritzi is abundant at all three limestone areas covered here and forms large colonies with bright green cordate leaves and the typical large, clavate appendix that is diagnostic for the species (Figures 17-18). Perhaps the most extraordinary species so far found, S. convolvula P.C.Boyce (Figure 19), is very locally abundant on the Padawan limestones. Homalomena that commonly occur include H. griffithii (Schott) Hook.f. (Figures 20-22) and H. insignis N.E.Br. (Figures 23-25). Homalomena and Schismatoglottis are frequently confused by non-specialists but are readily separable on inflorescence characteristics. In Schismatoglottis, the spathe limb is almost always shed during flowering (Figure 26), resulting in an infructescence that is fusiform with an open orifice (Figure 27). When ripe, the persistent lower spathe splits from the top, peeling back and reflexing to reveal the berries which, owing to the female flower zone being fused to the spathe, are held to one side (Figure 28). By comparison, the entire spathe persists into fruiting in Homalomena (Figure 29) and, at maturity the spathe splits from the base and curls upwards to reveal the cylindrical cluster of berries on their free stipe (Figures 30 & 31).
The rheophytic genera Bucephalandra and Piptospatha are closely related to Schismatoglottis and differentiated most readily by lacking a constriction at the junction of the lower and upper spathe, and a splash-cup infructescence, and from each other by spathe colour (white in Bucephalandra, pink in most Piptospatha), and the presence or absence of shield-shaped staminodes between the male and female flower zones –staminodes being present in Bucephalandra and absent in Piptospatha. Bucephalandra motleyana Schott is absent north of the Bongo Range and thus does not occur on the Bau limestones; however it is locally abundant on the Padawan and Serian limestones where it often forms large single-species colonies (Figures 32-34). Although Piptospatha does occur north of the Bongo Range, for example P. elongata (Engl.) N.E.Br. on the Lundu granites, there are no limestone-occurring species until south of the range where the recently described P. viridistigma P.C.Boyce, S.Y.Wong & Bogner is locally common on both the Padawan and Ranchan series (Figures 35 & 36). Cryptocoryne ferruginea Engl. is locally abundant in pools at the base of limestone cliffs in Bau, the leaves are often attractively bullate (Figures 37 & 38). The suffruticose genus Aglaonema is represented only by A. simplex (Figures 39 & 40) on exposed limestone, although A. nitidum (Jack) Kunth is frequently encountered nearby where the limestone is not exposed; A. simplex (Blume) Blume is not restricted to limestone and is a widespread and often very common species. These two species are frequently confused although they are readily separable by inflorescence characters, the most easily observed of which is that the spathe is caducous after anthesis in A. simplex but persistent through to almost full fruit maturity in A. nitidum (Figures 41-42). The climbing and hemiepiphytic aroids in Borneo are generally not narrowly endemic, and thus common and widespread species of Rhaphidophora and Scindapsus are found on limestones. Perhaps the commonest species on limestone is Rhaphidophora foraminifera (Engl.) Engl. (Figure 43), which is widespread through the Sunda Shelf, and with its adult leaves conspicuously perforated adjacent to the midrib and the leaf lamina abaxially yellow-pubescent (Figure 44), is readily identifiable. Another common species in the square-stemmed R. elliptica Ridl., most often encountered as a juvenile creeping along the forest floor on the lower parts of tree trunks, but occasionally seen as a many-stemmed hemiepiphyte reaching high into the canopy. These square-stemmed Rhaphidophora species are still very poorly understood, and it is increasingly likely that there are many species involved but as yet not formally recognized. Although most Rhaphidophora occurring on limestone are widespread species, there is at least one exception, R. tenuis Engl. (Figure 45), which while closely related to the most widespread species, R. korthalsii, is a limestone-obligate endemic to Borneo and easily identifiablei from R. korthalsii Schott by the shape of the juvenile shingle-stage leaves, the style of pinnation of the adult leaves, and by the slender spadix (Figures 46-48). Perhaps the most abundant Scindapsus in Borneo is S. longistipitatus Merr. (Figure 49), closely followed by S. treubii Engl., the latter with the juvenile stage often attractively variegated and the adult stage with markedly oblique leaf laminae (Figure 50). Less often seen on limestone is S. pictus Hassk. (Figure 51), a species well-established in commercial horticulture and also with often strikingly variegated juvenile plants but, like S. treubii, the adult plants lose their attractive variegation.
Pothos species on limestone, with the exception of the enormously variable P. scandens L. (Figure 52), tend to be restricted to that particular rock. Throughout Sarawak, Pothos ovatifolius Engl. (Figures 53-56) and Pothos insignis Engl. (Figures 57-59) are always encountered wherever limestone is present, even if the limestone is essentially almost completely underground, as on the southeastern flanks of the Bungo Range that divides Bau from Padawan. Pothos ovatifolius has a highly distinctive juvenile shingle-stage leaf lamina with the signature three intramarginal veins from the base crossing the primary lateral veins, while adult plants form extensive curtains of pendent stems flowering at the shoot tips, and later large infructescences. By contrast, P. insignis produces two distinct types of shoot. The clinging leaf shoots are sterile but give rise to clinging shoots clothed with large, somewhat inflated prophylls and cataphylls from between which arrive the inflorescences. These fertile clinging shoots can grow to several meters in length, producing a succession of several hundred inflorescences in flushes over a period of many years. Amydrium medium (Zoll. & Moritzi) Nicolson, the only species of this small genus in Borneo, is abundant on limestone as well as on virtually every other geology, and distinctive by the perforated and pinnate leaf lamina and white infructescences (Figures 60-62). Unlike virtually every other hemiepiphytic monsteroid, A. medium flowers only two or three metres from the ground. Anadendrum is perhaps the most poorly understood hemiepiphytic genus in Asia. Presently there are only eight published names but in Sarawak alone there appear to be at least 25 species. The limestone-associated species in west Sarawak (Figures 63-65) has the abaxial surface of the leaf lamina matte-glaucous grey, appears to be endemic, and is undescribed. Anadendrum is sometimes confused with Pothos but lacks the veins crossing the primary laterals and has truncate-topped fruits. We finish with a word of warning about working on limestone, especially the karst formations. The rocks weather to produce many knife-edge ribs and sink-holes, these often of some considerable depth and lined with more exceedingly sharp erosion ridges. Additionally the rocks are often extraordinarily slippery and so great care should be taken when clambering over such formations, and especially in testing before putting your weight onto a surface that while appearing solid may, in fact, consist only of a thick layer of leaf litter and humus supported on a few rotten fallen branches over a deep sinkhole. We have both acquired spectacular cuts while working on the limestones. | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Dr. Alfred Byrd Graf
1901-2001
Dr. Alfred B. Graf was born on the 23rd of November 1901 at the Weigelhof (Estate) in the suburbs of Nürnberg (Bavaria/Bayern, Germany) where his father was employed as a manager. On the 14th of December, 2001, Dr. Graf died peacefully in his sleep after a long illness in Düsseldorf (Nordheim-Westfalen, Germany); he was one hundred years old. His family had moved from Nürnberg to Weissenburg (Bayern) when he was a young boy and he spent most of his early years there. He immigrated to the United States from Germany and according to his diary arrived in New York by steamship on March 29, 1925. He immediately went by train to Sutton, Nebraska where he was obligated to work off his passage in exchange for farm labor. However in June, 1998, three years before his death, he returned to Düsseldorf in his native country where his wife, Liselotte (born Vorwerk) owned a home. Liselotte, who had been his lifelong helpmate and frequently traveled with him during his work, died there on the 22 of February, 2005. For most of his life, Graf and Liselotte lived in their house in East Rutherford, New Jersey, but some months of the year they spent in a second home in Vista, California. Dr. Graf had a large library in East Rutherford and in his workroom was a map of the world with a pin dot for every place he had visited. Due to his extensive travels, only a few areas remained uncovered with pins. I visited him once in East Rutherford and stayed overnight in his home.
During the first two years he lived in the United States, Dr. Graf lived in Nebraska and afterwards moved to Council Bluffs, Iowa where he started a small nursery and flower shop. However, he was driven out of business by a hail storm that destroyed his greenhouse. At that time, orchid growing was just beginning in the United States. Because he had already learned about tissue culture of orchids in his native Germany, this knowledge was in demand. Dr. Graf took a job at the Julius Roehrs Company, a large commercial nursery in New Jersey, where he spent the rest of his working career. There he became general operations manager for the company’s expanding greenhouse operation and it also became the base for his travels and research. The great interest shown by nursery customers for the company’s retail catalogue was the reason for Dr. Graf to broaden it into a book form. This was first published by him as, “Exotic House Plants Illustrated”, in 1953, with 178 pages and 1,200 illustrations. This book also appeared in several editions with the 10th edition appearing in 1976. The books allowed Dr. Graf to elaborate on the plants he had for sale and he added other pictures from plants he had photographed overseas. This first publication led him to later publish an enlarged book, “Exotic Plant Manual”, with 842 pages and 4,200 photographs, which also was published in several editions. The large book, “Exotica: Pictorial Cyclopedia of Indoor Plants”, (series 1) was published in December 1957. Several editions (all together 12 in four series) appeared and these were further enlarged, ultimately culminating in, “Exotica 4”, a work in two volumes with 2,606 pages and 16,300 photographs. Yet another book, “Tropica”, also went through several editions (fifth edition, 2003), with 1,154 pages (in the last edition) and now 7,000 colored photographs, listed virtually all cultivated plants originated from tropical and subtropical countries, It is still the biggest seller of Graf’s books. His last big book, “Hortica: Color Cyclopedia of Garden Flora in all Climates and Exotic Plants Indoors”, was published in 1992. It has 1,218 pages with 8,100 color photographs. All of these books were published over the years by the Julius Roehrs Company. Whereas the “Exotica” books contain mainly black and white photographs and only a few color plates, “Tropica” and “Hortica” are in full color. When Dr. Graf attempted to publish his first book, “Exotica”, and had approached commercial publishers, they thought it would be a risky venture for them to print the work. It was then that the Julius Roehrs Company decided to publish Graf’s books on their own and all of his publications became huge successes and with excellent and profitable sales.
Dr. Graf was a world authority on cultivated tropical and subtropical plants and was well known to gardeners, plant lovers and botanists all over the world. The pictorial cyclopedias of plants were of regular use by plant lovers who could determine their plants using photographs and find a key to their cultivation. As a globe-trotting botanist, plant hunter and collector, as well as a photographer, Dr. Graf searched throughout vast regions and remote islands. He visited the giant Lobelia on Mount Kilimanjaro in Africa, the Rhododendron in Sikkim (India) and the bamboos of China. He also brought back plants from his journeys. One of the most spectacular plants he introduced into cultivation was the Begonia exotica from New Guinea (it is correctly named Begonia brevirimosa Imsher today.) His travels also resulted in many plant photographs which were stored in his extensive archive. He was also in contact with many botanists and plant specialists around the world. When Alfred Graf came to Germany, he usually visited me at the Botanical Garden of München where I was employed for 33 years. We had long discussions about tropical and subtropical plants. Over the years, I sent him photographs of plants which we cultivated in our garden. Quite a number of them are included in his books. His correspondence to me was always in German and was broken only in his last years when he could not write any longer. By this time Dr. Graf’s declining eyesight made him totally blind and it was difficult for him to get around. The few remaining years prior to his death were tragic, especially since it was impossible for him to see the plants which were his life time’s work. Just a few months (since September, 2001) before his death, full time care was arranged for him and his wife who also grew less able to care for him. In 1962, when he traveled for 8 months through Africa, he caught an illness which was never diagnosed, even with the best doctors of tropical medicine in New York. The best they could do was to attempt to control the fever. The end result was damage to the retinas of both of his eyes. Even though he had made an incredible recovery, he had blank spots in his vision and much later he developed glaucoma and macular degeneration which probably was due to this earlier damage.
Dr. Graf received many honors during his life, the most important of which was the Dr. h.c. degree received in 1978 from Farleigh Dickinson University in Rutherford, New Jersey. This was a great moment in his life when his knowledge about plants was acclaimed by the academic community, especially since he was not trained at the university level and he was a self-made man in botany. Other honors were given to him by various horticultural organizations. Dr. Graf received the following awards: the Tercentenary Medallion from Gov. Rich Hughes of the State of New Jersey, the Citation Award from the American Horticultural Society, the Large Gold Medal from the New Jersey State Florists Association, the Large Gold Medal from the Massachusetts Horticultural Society in Boston, the Distinguished Service Award from the Horticultural Society of New York, the Certificate of Merit from the Pennsylvania Horticultural Society in Philadelphia, the Award of Merit and Silver Medal from the New York Florists Club, the Sarah Chapman Francis Medal from the Garden Clubs of America, induction into the Floriculture Hall of Fame by the Society of American Florists, the Liberty Hyde Bailey Medal from the American Horticultural Society and was given the Florida Hall of Fame Award by the Foliage Association in Orlando. Dr. Graf was also a contributor to “Encyclopedia Brittanica.”
“Exotica 4”, one of Albert Graf’s popular selling plant books, alone contains 1,248 aroids, many other plant families (813 Begoniaceae, 816 bromeliads, 1,021 cacti, 622 ferns, 533 gesneriads, 296 palms, 1,439 orchids and many others). The aroids are the second largest group of plants following the orchids. In addition to plant descriptions, a number of general chapters are included, such as how to care for plants indoors with a key to cultivation, hydroculture, gardening in warm climates, interior plantscaping, methods of propagation, insect and disease control, characteristics of plant families, botanical terms, pronunciation of Latinized plant names, history and origins of exotic plants, climate and vegetation maps, plant geography and habitats, and color charts used in horticulture and bibliography. There was also a “Synopsis of Revisions and Updates” in the 1985 edition. Dr. Graf was always very careful to get the correct name of a plant which was not so easy in many cases where plants were grown under a synonym or a completely wrong name.
I would like very much to thank Dr. Graf’s daughter, Doris Graf Matthews of Vista California , for information about her father for this obituary. My thanks also go to Jean Ruff from the Julius Roehrs Company in Farmingdale , New Jersey . Information was also obtained from his obituary in the New York Times.
Dr. Josef Bogner |