Rafflesia- The Worlds Largest Flower

Rafflesia- The Worlds Largest Flower

History

There is even more to tell of its ‘discovery’ and its claim to fame as the largest flower on earth. In the year 1818, Sir Stamford Raffles was posted as Governor to Bencoolen in Sumatra which was then, the administrative centre for the British East India Company for Western Sumatra. Raffles’s interest in natural sciences was insatiable.

A respected and popular member of the Royal Society in London, Raffles arranged and persuaded a fellow society member - Dr.Joseph Arnold to accompany him on an expedition into the interiors of Sumatra. It was on one of these expeditions that they stumbled on a discovery which was to puzzle botanists for a long time. Unfortunately, Dr Arnold died because of jungle fever before presenting the report to the society.

Habitat

Rafflesia are endemic to Southeast Asia recorded 17 known species, the Rafflesia flowers have been found only in Indonesia - Sumatra and Java, Malaysia, including Borneo island and south Thailand. Rafflesias have small, brownish, scale like leaves and fleshy, foul-smelling flowers of various sizes from few inches to meter big in diameter. Rafflesia classified as parasite, which means it just takes the nutrient out of its host.

The Rafflesia can be found at altitudes of between 500 and 700 meters in the forests of Malaysia, Southern Thailand, Sumatra and Java in Indonesia. In these tropical rainforests, the climate is continuously warm and humid, with very high humidity. The Rafflesia is rare and fairly hard to locate. It is especially difficult to see in bloom; the buds take up to 10 months to develop and the blossom lasts for just a few days. However, how many these strange plants are survived in primary rainforest is still unknown.

Characteristic

The cabbages like bugs that produce before blooming are size of cabbage. The large fleshy flower is what we usually notice; this is the flower’s sexual organ. There is a deep well in the centre of the flower containing a central raised disc that supports many vertical spines. The sexual organs are located beneath the rim of the disk, and male and female flowers are separate.

The reddish brown colors of the petals, sprinkled with white freckles exudes act most unpleasant stench, similar to rotting flesh or carrion. Some people believe that the stench attracts flies and insects which help disperse the seeds. Others believe that large animals could be agents for this seed dispersal. In order for the seed to germinate, it was found that the vine of the host plant must be damaged in some way so that the filaments of the seed may infiltrate successfully. The damage to the host vines could be made by trampling hoofs of large animals. The seeds adhere to the passing animals’ hoofs and are transported to other places where they can find host plants to attach to.

http://www.endemicguides.com/Rafflesia.htm

BUNGA RAFFLESIA

Wednesday, June 22, 2005

BUNGA RAFFLESIA

Photo by usman yasin

BUNGA RAFFLESIA

Rafflesia Arnoldii adalah salah satu jenis flora unik Indonesia yang dinobatkan sebagai "puspa langka nasional Indonesia". Ia mempunyai nama daerah yang beragam sesuai dengan bahasa penduduk kawasan tumbuhnya, seperti sekedai, ambun, bunga benalu, bunga hantu, ambai-ambai dan lain-lain. Ada beberapa macam bunga Rafflesia seperti Rafflesia Acehencis, Rafflesia Rochussenii, Rafflesia zollingeriana dan lain-lain yang tumbuhnya tersebar di beberapa daerah di kawasan Malenesia yang meliputi Malaysia, Indonesia dan Filipina. Tetapi jenis-jenis ini umumnya berukuran lebih kecil dengan penampilan saling berbeda. Rafflesia Arnoldii berukuran raksasa dan diketahui hanya terdapat di Sumatera dan penyebarannya berada di sepanjang punggung Bukit Barisan dari Aceh sampai Lampung dengan pusat ekologi di Bengkulu

Pertumbuhan Rafflesia Rrnoldii dimulai dengan perkecambahan yang terdapat di dalam kulit tumbuhan inang kemudian berkembang menjadi benang-benang

Proses terbentuknya bunga diawali oleh pembengkakan di dalam akar atau batang tumbuhan inang serta terbentuknya kuncup. Kuncup ini terus membesar sampai secara perlahan merobek permukaan. Kulit inang pecah sehingga terlihat bagian kuncup yang diliputi oleh braktea berwarna putih yang kemudian berubah menjadi coklat kehitaman. Pada diameter sekitar 25 cm, braktea tergeser dan terlepas satu persatu sehingga terlihat bagian bunga berwarna merah muda, bagian ini merupakan bagian yang kelak menjadi perigonium ( perhiasan bunga ). Braktea dapat dibedakan dari perigonium yaitu dari warnanya yang lebih gelap, lebih keras dan lebih tipis. Bunga mulai mekar dengan membukanya lobur perigonium satu persatu atau kira-kira pada saat kuncup berdiameter 30-35 cm. Lamanya perkembangan dari kuncup yang berdiameter 4 cm sampai bunga mekar ( diameter kuncup sekitar 34 cm ) diperkirakan 310 hari. Sedangkan waktu yang diperlukan dari fase biji sampai terbentuknya biji lagi diperkirakan selama 4,5 – 5 tahun.

Masa mekar sampai layu bunga Rafflesia Arnoldii biasanya 5-7 hari, kemudian membusuk dan biasanya akan dikerumuni lalat dan serangga lain. Rafflesia Arnoldii berbunga sepanjang tahun dan saat berbunga paling banyak adalah pada bulan-bulan basah.

Saat mekar, bunga Rafflesia Arnoldi mengeluarkan bau agak busuk. Sehingga ada yang menyamakan namanya dengan bunga bangkai ( Amorphophallus titanum ). Selain itu Rafflesia Arnoldii juga dikenal dengan sebutan “Padma Raksasa” karena ukurannya yang besar.

Bau busuk dari Rafflesia Arnoldii akan menarik berbagai jenis serangga terutama lalat. Lalat ini akan hinggap dari satu bunga ke bunga yang lain. Rafflesia Arnoldii merupakan tumbuhan berumah dua, sehingga dalam penyerbukannya memerlukan perantara yang berupa hewan. Lalat merupakan hewan utama yang membantu dalam penyerbukan. Lalat penyerbuk pada tumbuhan ini adalah Lucilia sp (lalat hijau) dan Sarchopaga ( lalat abu-abu ). Jika bunga betina dapat diserbuki maka akan dihasilkan buah yang berisi lebih dari 100 biji. Bunga jantan dan bunga betina akan sulit dibedakan apabila kita lihat dari luar karena kedua-duanya berwarna merah kecoklat-coklatan dengan bintik-bintik putih.

Biji Rafflesia Arnoldii yang terdapat pada jaringan buah yang terurai hanya dapat tumbuh pada tumbuhan inangnya bila terdapat hewan penyebar biji yang berfungsi sebagai pembawa biji dan melukai akar tumbuhan inang. Hewan yang berperanan dalam penyebaran biji ini diduga berasal dari mamalia berkuku ( ungulata ) seperti babi hutan, rusa, kijang dan jenis tupai.

Diposting oleh Usman Yassin

http://yayasan-lembak.blogspot.com/2005/06/bunga-rafflesia.html

Rafflesia Arnoldii : Masuki Musim Mekar di Bengkulu

Masuki Musim Mekar di Bengkulu

Jan 3, '08 12:30 AM for everyone

Category:

Other

Rafflesia Arnoldi

Menjelang akhir tahun 2007 bunga langka Rafflesia Arnoldi mulai memasuki musim mekar. Sejak sepekan belakangan setidaknya ditemukan lima kuntum bunga tengah mekar di kawasan hutan lindung Register 5 Bukit Daun, Taba Penanjung, Bengkulu Utara, sekitar 52 km sebelah timur Kota Bengkulu.

Lokasi mekarnya bunga Rafflesia ini seperti yang disaksikan Kompas hari Minggu (30/12), berada di tiga lokasi secara terpisah di kawasan hutan lindung Bukit Daun. Menariknya, tempat mekar bunga ini ternyata berada persis di pinggir jalan nasional Bengkulu – Kepahiang – Curup - Lubuk Linggau. Sejumlah wisatawan lokal baik asal Bengkulu maupun dari Lubuk Linggau dan Pagar Alam, Sumatera Selatan menyaksikan mekarnya bunga-bunga itu.

Menurut Rizal dan Antoni, dua warga Taba Penanjung yang menjaga bunga-bunga Rafflesia tersebut, masa puncak mekarnya bunga langka ini memang berlangsung pada bulan Desember. Pada bulan itu, bonggol bunga yang mirip dengan kol ini akan muncul dari inangnya dan mekar dalam tujuh hari. Setelahnya, bunga yang menjadikan serangga kecil sebagai santapan itu akan layu.

“Hutan Lindung Bukit Daun yang dibelah jalan nasional Bengkulu – Curup menuju Lintas Tengah Sumatera ini, memang menjadi tempat tumbuh alami sejak lama," kata Rizal. "Jika ditelusuri sampai ke bagian dalam kawasan hutan lindung, pasti banyak ditemukan bunga yang tengah mekar. Namun, agar masyarakat bisa menyaksikan keindahannya maka kami berupaya mencari lokasi yang dekat dengan jalan raya,” tutur Rizal.

Padma raksasa Rafflesia Arnoldi, pertama kali ditemukan oleh Thomas Stanford Raffles dan ahli botani Arnol pada tahun 1818 di kawasan hutan di Manna, wilayah Bengkulu Selatan sekarang. Belakangan sejumlah kawasan hutan di Provinsi Bengkulu menjadi habitat alami tempat tumbuhnya bunga Rafflesia tersebut. Bunga yang berukuran besar itu kini menjadi maskot pengembangan ekowisata di Provinsi Bengkulu.

Achmad Zulkani
Copyright Kompas

Parasitic plant research leads to article

Parasitic plant research leads to article

SIUC professor gets published in Science magazine

BY CALEB HALE, THE SOUTHERN

Wednesday, January 17, 2007 7:08 AM CST

CARBONDALE - The sheer size of it evokes bewilderment and slight terror. Its three-foot diameter is an overload to the human senses, and its bizarre blood red and white-speckled petals appear other-worldly compared to other flora.

What mutes these effects is the fact this particular Rafflesia arnoldii is a cardboard cutout of the real thing - a Halloween costume actually, says Southern Illinois University Carbondale plant biologist Daniel Nickrent, demonstrating its use by putting it over his head with a big grin.

The real Rafflesia arnoldii, a parasitic flower native to southeastern Asia, can grow to become a 15-pound monstrosity, with a gaping maw at its center and capable of producing a scent somewhat like rotting flesh.

Nickrent has spent the last 16 years at SIUC studying parasitic plants such as Rafflesia, despite their odd quirks. Now, thanks to recently co-authoring an article with Harvard professor Charles Davis, Nickrent expects to see his research in print in the pages of Science magazine soon.

The article, Nickrent's first in Science, appeared Jan. 11 in the publication's online version. The selection of the article is prestigious, he said; he hopes it will be remembered by someone when it is time to submit grant proposals later this year.

"(Science) is extremely widely circulated and read," Nickrent said. "It's broad. I call it a magazine versus a journal because it tends to touch on a wide range of subjects of general interest. And, of course, they want to sell, so they can pick and choose what they want to feature."

The article points out a revelation in the world of plant science. It identifies that Rafflesia has common origins with the same group of plants that produce more common flowers, such as poinsettias and violets.

Nickrent said tracing Rafflesia's roots has always been difficult because it essentially lacks all of the key identifiers biologists use to classify flowers.

"The Rafflesia group has been one that's intrigued everybody; nobody knew quite where it belonged," Nickrent said.

The first cited discovery of Rafflesia came in 1818. The plant was named after Sir Thomas Stamford Raffle, who was governor of Singapore at the time under British rule.

Rafflesia is only found in Southeast Asia, in the jungles of Thailand, Indonesia and Malaysia, for instance. Even there, Nickrent, who has visited the area, said finding one of the flowers in full bloom doesn't happen that often.

"These are some of the rarer plants in the world. To find one in flower is a real treat," he said.

Southeast Asians believe the plant has medicinal value for postpartum women; hence some of the markets in the region are filled with crated Rafflesia buds. Nickrent said that trade could someday place the flower on an endangered list.

Nickrent plans to continue his research into Rafflesia, particularly into the genomes that make the plant the way it is. He is co-authoring a forthcoming book with Old Dominion University professor Lytton Musselman, "Parasitic Plants of the World."

In the meantime, Nickrent continues to chronicle his research on his Web site, "The Parasitic Plant Connection," accessible from the SIUC Web site.

Interim College of Science Dean Jim Tyrrell said the national recognition a published article in Science brings is a feather in the college's cap.

"That is a very positive step. Science is, without question, the leading publication of the specialized disciplines," Tyrrell said. "There are other people in the college who have published in Science �¿� but this is not the usual run-of-the-mill science project; it is a bit distinctive."

Tyrrell said the College of Science has been doing well in recent years with external federal research funding. The college had a portfolio of $5.5 million in the 2001 fiscal year, and by last fiscal year the amount had risen to roughly $9 million.



caleb.hale@thesouthern.com

(618) 529-5454 ext. 5090

h*tp://www.thesouthern.com/articles/2007/01/17/local/18895681.txt

Rafflesia (Rafflesia kerrii), Khao Sok National Park, Thailand

Rafflesia (Rafflesia kerrii) flower and bud, Khao Sok National Park, Thailand

Location: Thailand, Muang Thai ; Indochina ; Southeast Asia ; Asia ;

Keywords: parasitic, parasite, parasitic plants, Tetrastigma, Vitaceae, Liana, Rafflesia kerrii ; Rafflesia ; Rafflesiaceae ; Malpighiales ; Rosidae ; Eudicots, Rosopsida ; Magnoliophyta, flowering plant ; Angiospermae, angiosperm ; Spermatophyta, spermatophyte ; Vascular plant, Tracheobionta ; Embryophyta ; Streptophyta ; Viridiplantae ; flora, Plantae, plant ; bloom, flower; flower bud, floral bud; flowering, blossoming, blossom, blooming; vegetation; parasite; monocarpic; rainforest; National Park;

Date: 14.08.2007
[M]: not applicable
Category: Nature
colour space: RGB
size in px: 3008 x 1996
Type: photo
Colour: colour
Format: horizontal
Original available as: tif file (or similar, uncompressed)
Scan available as: not applicable
Exclusive rights: Exclusive usage rights available on request
Licence: Simple Licence
Author:

Bernd Mehmen
Ringstrasse 20
42349 Wuppertal
Deutschland

b.mehmen@freenet.de

Rafflesia kerrii Meijer, Thailand

Rafflesia kerrii Meijer

Species named after A. F. G. Kerr, botanist employed by the government of Thailand.

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A male flower of R. kerrii measuring 0.5 m in diameter. Two blowflies can be seen on the right perigone lobe and two are in flight above the diaphragm on the left. Prachuab Khirikhan, Thailand. Photograph by H. Bänziger, from Bänziger (1991) "Stench and fragrance: unique pollination lure of Thailand's largest flower, Rafflesia kerii Meijer" (Nat. Hist. Bull. Siam Soc. 39:19-52).

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Rafflesia kerrii, open flower, approximately 80 cm (36 inches) in diameter. Rainforest in the Lojing Highlands (near the Cameron Highlands), Kelanatan, Malaysia. Photo by Fletcher & Baylis, from Wildside’s Searchable Album.

Rafflesia kerrii. Flower next to guide [Orang asli, the indigenous people of the area]. Flower is approximately 80 cm (36 inches) in diameter. Rainforest in the Lojing Highlands (near the Cameron Highlands), Kelanatan, Malaysia. Photo by W. K. Fletcher & D. M. Baylis, from Wildside’s Searchable Album.

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Open flower. Khaosok rainforest, northern end of southern Thailand. Photo taken January 23, 2001 by Ray Merrell.

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SIUC / College of Science / Parasitic Plant Connection / Rafflesiaceae
URL: http://www.parasiticplants.siu.edu/Rafflesiaceae/Raff.kerrii.page.html
Last updated: 19-Oct-05 / dln

http://www.parasiticplants.siu.edu/Rafflesiaceae/Raff.kerrii.page.html

World's Second Largest Flower Thought Extinct 'Rediscovered' in Region X

By Mike Baños



In an event similar to the discovery of the "extinct" coelacanth off the coast of South Africa in 1938, a species considered the world's second largest flower previously though to be extinct has been found in a forested area of Northern Mindanao.

Two weeks ago, reports of a strange flower blooming in the mountains of Region 10 reached the Protected Areas and Wildlife Division, Region 10 office (PAWD-10). A team dispatched by Director Malou Clarete verified that the large bloom was indeed that of the Rafflesia schadenbergiana Greppert, named after German botanist Alex Schadenberg who discovered it in Mt. Apo with fellow German Otto Koch in 1882 (The Snows of Mt. Apo, from History Against the Landscape by Miguel A. Bernad, S.J., 1968).

It was the first and last time people would see the giant flower for the next 125 years. At 80 centimeters across, it is considered to be the second largest flower in the world. According to Fr. Bernad's account of the Schadenberg expedition, "We are told that young buds of this plant, growing together on one stem, were found on one occasion to weigh as heavily as a double-barreled gun and six solid bullets. (Bernad, 1968).

The latest sighting is not just of one flower but rather what PAWD-10 calls a "population." It is the only known population existing in the Philippines of R. schadenbergiana that is actively flowering.

The cluster was found in a community-based forestry management (CBFM) area and includes six more flower buds which have not yet bloomed. It could take from 9-10 months after they first appear for buds to bloom. Not all buds bloom into flowers, with many decaying unopened. Too much rain can cause buds to rot, too little cause them to shrivel and dry up.

The first flower which blossomed was measured at 70 centimeters across and has been taken by Dr. Julie Barcelona of the Philippine National Museum to Manila for preservation. Together with the PAWD-10, the National Museum has sat down with the local government where the R. schadenbergiana population is found, the Department of Tourism, DENR and the academe to develop a protection and management plan for the "resurrected species."

"We can still not divulge the exact location of the population at this time pending the declaration of the area as a critical habitat by Environment Sec. Angelo Reyes," Ms. Clarete said. "DENR is still conducting a site assessment of the area which a requisite of its declaration as a "critical habitat" for the specimens.

"We fear people might stampede to the area and adversely affect the fragile flowers, their buds or host-vines," said tourism director Catalino Chan III. "We first want to establish the carrying capacity of the area before we allow visitors, considering the specie in question is considered critically endangered."



Section 27 of Republic Act 9147 prohibits the collection, possession, transport and trading of all Rafflesia species listed as a critically endangered species under DENR Administrative Order #2007-01 on pain of 6-12 years imprisonment or a P100, 000-1,000,000 fine.

The PAWD says the Philippines has the highest density of Rafflesia in the world relative to land area. Unfortunately, the lowland rainforest it calls home is under siege from deforestation and conversion to other uses.

The first species identified as Rafflesia manillana Teschem was found in Basey, Samar in 1841. Other species subsequently discovered in the Philippines include R. cumingi (1845), R. philippensis Blanco (1845) and R.lagascae Blanco (1879, now both considered the same as R. manillana), and more recently, R.baletei Barcelona & Cajano, Camarines Sur; R. lobata Galang & Madulid, Panay Island; R. speciosa Fernando & Ong (2002) Sibalom, Antique, and R. mira. Fernando and Madulid (2005) Mt. Candalaga, Compostela Valley. All species are endangered or threatened. Rafflesia species are classified in the division Magnoliophyta, class Magnoliopsida, order Rafflesiales, family Rafflesiaceae.

What could be the smallest Rafflesia in the Philippines (measuring 12-13 cm in diameter) was recently discovered in Mt. Asog in Camarines Sur,. The researchers from the Camarines Sur State Agricultural College who discovered it has proposed the species be named Rafflesia irigaenses.

According to wikipedia, the free online encyclopedia, Rafflesia is a genus of parasitic flowering plants discovered in Indonesia by an Indonesian guide working for Dr. Joseph Arnold in 1818, and named after Sir Thomas Stamford Raffles, leader of the expedition. It has some 20 recognized species, all found in southeastern Asia, on the Malay Peninsula, Borneo, Sumatra and Kalimantan, West Malaysia, and the Philippines. Many of the species are extremely rare, and have been recorded in only a handful of localities.

The plant has no stems, leaves or true roots. The flower is a parasite which grows within its host, the tetrastigma vine, and appears as a tangle of fibers in its early stages. It only starts manifesting itself during its reproductive cycle. Outgrowths appear on the root vine, which develop into cabbage-like buds, followed by the blooming of a fully open flower which later bears fruit.

Its largest species, the Rafflesia arnoldii found in Malaysia, has a flower over a meter in diameter and weighs up to 10 kilograms. Even its smallest recognized species, R. manillana, has 20 cm. diameter flowers.

The survival of the sensitive Rafflesia depends on a lot of factors—its seeds need to find the right host, the buds need to receive the right amount of water and nutrients, and flowers of the opposite sex must be nearby so that pollination can occur.

Park authorities in Malaysian natural parks where the flowers have been star attractions for years have instituted measures to minimize the impact of tourism on the survival of the flowers. A lot of buds reportedly failed to bloom when disturbed. Cultivating the flower has met little success.

The PAWD says the most significant threat to the Rafflesia is the destruction of its rainforest habitat. Every year, 100,000 hectares of rainforests across the country are being felled due to logging, mining, and the conversion of forest lands for commercial and residential uses.

http://www.iligan.gov.ph/forum/index.php?topic=64.msg790

Rafflesia Speciosa : Sibalom NaturePark,Antique, Panay,Philippines

Rafflesia Speciosa

Sibalom NaturePark
Antique, Panay
Philippines
Mar. 5 2005

h*tp://www.mytropicalvisions.com/animality/rafflesias.html

Rafflesia (Rafflesia spp)

Opening bloom of Rafflesia speciosa and a dead bloom in the foreground in rainforest

http://www.arkive.org/species/GES/plants_and_algae/Rafflesia_spp/GES025158.html

Rafflesia Priceii : Sabah,Malaysian Borneo

Tambunan Rafflesia Conservation Area
Sabah,Malaysian Borneo
January. 13, 2003

h*tp://www.mytropicalvisions.com/animality/rafflesiaspeciosa.html

Bunga Rafflesia

Sayangi Bumi

Bunga Rafflesia

Bunga Rafflesia atau juga dikenali sebagai bunga pakma adalah sejenis tumbuhan yang unik. Ia tidak mempunyai daun, batang dan akar. Ia tumbuh di atas kayu tertentu seperti kayu vines (Tetrastigma), yang selalu didapati di dalam hutan tropika di Asia Tenggara. Ia tidak seperti tumbuhan berbunga lain kerana kebanyakan hidupnya ia sebagai bebenang nipis yang membuat linkaran di dalam kayu bagi menyebarkan benihnya. Ia seakan-akan fungus tetapi mangeluarkan bunga yang mempunyai stamens dan pistils.

Bunga Rafflesia hanya boleh dijumpai apabila ia mula untuk berkembang. Ia kelihatan seperti kelopak kubis. Banyak kuntum bunga yang busuk dahulu sebelum ia sempat kembang. Tetapi selepas 9 bulan, ia akan mula berkembang sehingga berdiameter 1 meter dan seberat 10kg.

Ia berbau seperti cendawan. Ditengah-tengah bunga terdapat objek yang tumbuh yang tidak diketahui apa kegunaannya. Ia mengandungi banyak nectar, ini adalah untuk menarik perhatian dan memerangkap serangga kecil yang hinggap diatasnya. Selepas 3-4 hari, bunga Rafflesia mula mengecut dan warnanya bertukar menjadi warna hitam.

Bunga-bunga ini terdiri daripada jantan dan betina. Bunga betina jarang dijumpai. Bunga pakma juga menghasilkan buah. Buahnya berbentuk bulat dan bersaiz 15 cm. kulitnya berminyak dan dipenuhi biji benih. Isinya dapat menarik perhatian tupai dan tikus pokok yang selalunya menjadi agen menyebaran bagi benih-benih refflesia.

Bunga Rafflesia bukan saja merupakan bunga terbesar malah terberat. Berat bunga refflesia adalah 7kg. Tumbuhan ini dijumpai oleh Stamford Raffles. Raffles menjumpai bunga rafflesia di Borneo, Sumatra dan jawa.

Terdapat 14 spesis bunga Rafflesia, Rafflesia arnoldii merupakan yang terbesar sekali. Usaha untuk menanam bunga Rafflesia di taman botani adalah sukar.

Kita patut berbangga kerana negara kita merupakan salah satu habitat bunga Rafflesia. Oleh itu peliharalah ia semoga generasi akan datang dapat menikmati keindahan semula jadi yang tidak terperi ini.

	Rafflesia micropylora Meijer North Sumatra (Indonesia)
	Rafflesia arnoldii R. Brown Borneo (Sarawak, Malaysia) West Kalimantan (Indonesia) Benkulu, Sumatra (Indonesia)
	Rafflesia cantleyi Solms-Laubach Peninsular Malaysian & Tioman Island
	Rafflesia pricei Meijer Borneo (Brunei, Sabah, N. Sarawak ?, Malaysia) Kalimantan? (Indonesia)
	Rafflesia tuan-mudae Becc Form restricted to three isolated Mts. in W. Sarawak (Malaysia). Similar to R. arnoldii.
	Rafflesia gadutensis Meijer Western coastal Sumatra and Benkulu (Indonesia)
	Rafflesia kerrii Meijer Peninsular Thailand and Malaysia
	Rafflesia speciosa Island of Panay (Philippines)
	Rafflesia keithii Meijer Borneo (Sabah, Malaysia) East Kalimantan (Indonesia)
	Rafflesia speciosa Island of Panay (Philippines)
	Rafflesia manillana Teschemacher Luzon (Philippines)

http://www.epgenius.com.my/H_000007.asp

Rafflesia Patma

Rafflesia patma mempunyai ukuran diameter bunga bisa mencapai 25 cm lebih. Organ penyusun bunga dari Rafflesia patma tidak jauh dari Rafflesia arnoldii namun ukurannya lebih kecil. Sosok Rafflesia patma sepintas memang mirip dengan Rafflesia yang lain, hanya saja bila diamati lebih teliti mahkota bunganya memiliki warna yang lebih pucat. Saat mekar sempurna diameter bunganya berkisar 20 – 30 cm. Tanda yang khas, yang membedakan dengan Rafflesia rochussenii adalah duri – duri yang terdapat pada diskus.

Rafflesia patma selain tumbuh pada akar inangnya juga tumbuh pada batang inang yang menggantung di atas lantai hutan, tetapi persentase yang tumbuh pada bagian bukan akar sangat kecil dan tumbunya tidak sebaik yang tumbuh di akar inang. Diameter akar inang yang baik untuk ditumbuhi Rafflesia patma berkisar antara 1,1 – 3 cm. Ukuran kuncup Rafflesia patma yang ditemukan di Cagar Alam Leuweung Sancang sangat beragam mulai dari diameter 2 cm sampai 15 cm. Persentase kematian kuncup yang paling banyak terjadi pada diameter <>et al, 1989 ).

Kondisi ekologis Rafflesia patma yang terdapat di Cagar Alam Leuweung Sancang Jawa Barat adalah sebagai berikut : Jenis tanah regosol, kelas tekstur tanah lempung berpasir, konsistensi tanah gembur dengan kelas drainase baik, pH tanah agak masam sampai netral; kandungan C organisk dan Ca sangat tinggi; N total, Mg dan kapasitas tukat kation (KTK) tinggi; P tersedia sangat rendah; K dan Na sedang dan tumbuh pada ketinggian berkisar antara 0 – 35 m dpl dengan jarak dari pantai berkisar antara 5 – 700 m dan kemiringan lahan tidak lebih dari 5 %. Penyerbukan terjadi diduga dengan bantuan lalat, yang mendatangi bagian reoriduksi bunga adalah lalat hijau (Lucillia sp.), lalat biru (Protocalliphora sp). Lalat abu–abu (Sarcophaga sp) dan lalat buah (Drosophila sp). Sedangkan penyebaran biji diduga dilakukan oleh rayap tanah (Macrotermes sp), semut merah besar (Polyergus sp), babi hutan (Sus scroffa Linn), landak (Hystrix brachyura Linn), tupai (Tupaia sp) dan muncak (Muntiacus muntjak Zimm) (Priatna et al., 1989).

Habitat yang sesuai untuk Rafflesia patma adalah daerah antara tipe lautan pantai dengan tipe hutan hujan tropika dataran rendah (ekoton). Asosiasi tingkat pohon terdiri dari : Aleurites moluccana, Tabarmaemontana sphaerocarpa, Neesia altissima, Pongamia pinnata, Dracontomelon mangiferum, Cinnamomun inners, Tarenna incerta, Barringtonia acutangula, Terminalia catappa. Asosiasi vegetasi tingkat belta adalah Tetrastigma sp, Calamus ornatus, Cinnamomun inners, Pterospermum diversifolium dan Dracontomelon mangiferum. Sedangkan asosiasi tingkat semai dan tumbuhan bawah adalah Colomus orantanus, Pipturus repandus dan Tarenna incerta. Penyebaran Rafflesia patma di Cagar Alam Leuweung Sancang secara sporadis meliputi areal yang berdekatan dengan pantai pada tinggi berkisar antara 0 – 35 m dpl dan jarak dari pantai berkisar antara 5 – 700 m kemiringan lahan umumnya diatas atau tidak lebih dari 5 %

Acaman kelestarian yang terjadi pada Rafflesia patma yang terdapat di Cagar Alam Leuweung Sancang adalah terjadinya pemungutan kuncup oleh masyarakat untuk bahan baku jamu (Priatna et al, 1980)

http://www.rafflesia.info/patma.htm

Is Rafflesia lobata real?

01 April 2007

Is Rafflesia lobata real?

When Dr Domingo Madulid sent me the paper on the new species of Rafflesia from Panay to be published in Folia, I was very sceptical. In fact I was sceptical until yesterday! The reason was simple: there was very little features in this new species, called R. lobata from R. manilana. I blog about this in December last year, here.

However, my good friend and companion, Dr K Fletcher of Wildsidephotography returned from Panay Is last week with splendid photographs of R. lobata in the wild. I was given a copy of his photos and reproduced here.

I have no doubt after viewing these photos that those lobbed diaphragm is an artifact. The colors of the the diaphragm and the disc are also not the same as R. manilana. I can confirm now that this species is a good one. Despite my initial sceptism, I am now comvinced that this species is in no way the same to R manillana. Fletcher also reported that the R. lobata grows on limestone hill, which is not a case for R. manilana.

I got to visit this place !

Posted by Prof Dr Kamarudin Mat-Salleh at 1:08 AM
ht*p://rafflesia-in-bloom.blogspot.com/2007/04/is-rafflesia-lobata-real.htm

Rare plant species found in Mt. Kanlaon

Cebu Daily News
First Posted 12:07pm (Mla time) 01/25/2008

BACOLOD CITY—A research firm commissioned by the Philippine National Oil Co.-Energy Development Corp. (PNOC-EDC) found a rare parasitic flowering plant in the Mt. Kanlaon Natural Park.

The biodiversity study undertaken by Maunsell Philippine Inc. that was released on Tuesday noted that it found Rafflesia speciosa, a member of the family of the world's largest flower, in Mt. Kanlaon Natural Park's 169-hectare buffer zone.

"The species is the first record of this unique genus in the island (of Negros). The genus Rafflesia is very rare and can only be seen in the forests of Panay, Palawan, Makiling and Banahaw," said the PNOC-EDC commissioned Maunsell Philippines Inc. Biodiversity Study furnished to the INQUIRER.

Among the seven recorded Rafflesia species in the country, the Rafflesia speciosa has the largest flower that could attain a maximum diameter of two feet, the PNOC-EDC study said. The one found in the MKNP buffer zone was about 30 centimeters in diameter. /Inquirer

ht*p://globalnation.inquirer.net/cebudailynews/visayas/view_article.php?article_id=114732

Observatory

A Smelly Puzzle, Solved

Courtesy of Jeremy Holden

QUITE A BOUQUET Rafflesia arnoldii is the world’s largest flower and smells like rotting flesh. It is also a parasite and lacks roots and leaves.

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By HENRY FOUNTAIN

Published: January 16, 2007

Rafflesia arnoldii is no shrinking violet. At up to three feet in diameter, it’s the world’s largest flower. It’s also possibly the most repulsive — it looks and smells like rotting flesh (the better to attract flies, which act as pollinators).

But while Rafflesia may be easy to describe, it has been much harder to classify. It’s a parasite, embedding itself in vines in the understory of pristine rainforests in Indonesia, and it lacks the roots, stems, leaves and photosynthesizing machinery that would give scientists a clue as to its evolutionary background.

Rafflesia and related species “really have remained a mystery,” said Charles C. Davis, an evolutionary biologist at Harvard. Genetic analysis is required, and although initial work by Dr. Davis and others put them in the order Malpighiales, the research was insufficient to pinpoint their place within the order.

Now, Dr. Davis and colleagues have done more genetic research to solve the puzzle. Their conclusion, published online by the journal Science, is that the Rafflesiaceae, as this family of species is known, are nestled within the spurge family, which includes rubber plants, castor, cassava and poinsettia. “They are smack dab in the middle,” Dr. Davis said.

In some ways this is a surprise, because spurges are so well known. On the other hand, Dr. Davis said, the Rafflesiaceae “are so off on their own trip that their position within any group would require some explaining.”

One thing that requires explaining, Dr. Davis said, is the remarkable range in size. Among the spurges are some with very small flowers. The researchers estimated that as the Rafflesiaceae diverged over 46 million years, floral size increased 79-fold.

“These plants flower in understory rainforest environments, which are dimly lit, so they are not easily seen by pollinators,” Dr. Davis said. “They would have had incredible incentive to increase their surface area, to maximize odor production and bring in these pollinators.”

Same Lobsters, More Whales

Maine lobstermen work extremely hard, baiting, setting, hauling and repairing traps just about every day of the year.

But do they work too hard? Ransom A. Myers of Dalhousie University in Nova Scotia and colleagues argue that they do, and that by slacking off a bit they could help protect the North Atlantic right whale.

The whales have been in dire straits for decades, and currently number about 350 individuals off the coast of North America. Although they are a protected species, accidental deaths have slowed their recovery. Major causes of death are collisions with ships and entanglement in fishing gear, including the lines connecting lobster traps to surface buoys and to one another.

In a study in Current Biology, the researchers compared lobstering on the American and Canadian sides of the Gulf of Maine. On the Canadian side, southwest of Nova Scotia, lobstering is allowed from December to May, and fewer than 400,000 traps are used. In Maine, lobstering is an all-year activity, with 3.2 million traps.

Yet for all those differences, the Maine harvest is just 30 percent greater than on the Canadian side. The researchers estimate that Maine lobstermen could cut their season in half, reduce the number of traps by a factor of 10 and still harvest as many lobsters.

Most sightings of right whales in the Gulf of Maine occur in spring and summer. By shifting the Maine lobstering season and greatly reducing the number of traps, the researchers say, the risk to the whales would be greatly reduced.

Moths and a Drink of Tears

Moths and butterflies obtain moisture wherever they can find it — in Africa, Asia and South America, even from the tears of mammals and reptiles. But until now, no moth or butterfly has been seen drinking tears from a bird.

Roland Hilgartner of the University of Ulm in Germany and colleagues observed a species of moth in Madagascar, Hemiceratoides hieroglyphica, that alights on the neck of a sleeping magpie or Newtonia bird and sticks its long proboscis between the bird’s closed eyelids. Moths were observed in this position for 30 minutes or longer, presumably drinking the bird’s tears. The finding was reported in Biology Letters.

The moth’s proboscis is about half an inch long (about half the moth’s length), with a sharp point and many tiny spines and barbs. The researchers suggest that it functions somewhat like a harpoon, because it has to go not only between upper and lower eyelids, but also through the bird’s nictitating membrane, which further protects the eyes.

In this way it is more like bloodsucking moths and less like other tear drinkers, which generally have soft-tipped proboscises.

http://www.people.fas.harvard.edu/~ccdavis/weblinks/New_York_Times.htm

World's Largest Flower Mystery Solved

mercredi 14 janvier 2004

World's Largest Flower Mystery Solved The world's largest flower, called Rafflesia, can have a diameter up to one meter and can weigh up to 10 kilograms. It also smells like rotting flesh. Discovery News tells us that its genetic roots have been uncovered and that this plant that smells so bad is related to delicate flowers such as poinsettias or violets. The flower, which is one of 20 species collectively called Rafflesia, is related to poinsettias, violets, passionflowers, and other members of the order Malpighiales, according to a paper published in the current Proceedings of the National Academy of Sciences (PNAS). Researchers also have some intriguing theories as to why Rafflesia smells so awful, why its flowers bloom only once a year and live for five to seven days, why the plant is a parasite, and why it likes to attract flies that normally go for mounds of dead flesh. Before going further, here is a photo showing a man surrounding a rafflesia (Credit: Bruce Coleman Collection, photo by Allain Compost). How did the researchers proceed? Rafflesia is a parasitic plant that lacks roots, stems, and leaves. Usually botanists trace plant orders using chloroplast DNA, but since Rafflesia does not have chloroplast genes, Todd Barkman, assistant professor of biological sciences at Western Michigan University, and his team instead analyzed the Rafflesia's mitochondrial DNA (mtDNA), which corresponds to chloroplast DNA. The mtDNA studies revealed that the large, stinky flower is related to more normal-sized, pleasantly fragrant posies. Despite its enormity, Rafflesia does resemble passionflowers, according to Barkman. Both have their stamens and pistils fused together in a central column, and both produce a corona, or crown, in the shape of a ring. Here is another photo showing details of this flower (Credit: Todd Barkman). Rafflesia did not begin its life as a parasite, but likely evolved this lifestyle in Southeast Asia as it began to depend upon a variety of grape, Tetrastigma. Now Rafflesia lives inside of the grape plant and only reveals itself once a year when it produces the smelly blooms. "By human standards, the large, lumpy, mottled, stinky flowers could be interpreted as mimicking rotting flesh," Barkman told Discovery News. "The fact that flies are highly attracted to the flowers suggests that they think the flowers look and smell like rotting flesh too!" For more information about the rafflesia, you can look at this definition on Wikipedia and at this Rafflesia gallery. Finally, here is the abstract of the paper published by PNAS, "Mitochondrial DNA sequences reveal the photosynthetic relatives of Rafflesia, the world�s largest flower." All parasites are thought to have evolved from free-living ancestors. However, the ancestral conditions facilitating the shift to parasitism are unclear, particularly in plants because the phylogenetic position of many parasites is unknown. This is especially true for Rafflesia, an endophytic holoparasite that produces the largest flowers in the world and has defied confident phylogenetic placement since its discovery >180 years ago. Here we present results of a phylogenetic analysis of 95 species of seed plants designed to infer the position of Rafflesia in an evolutionary context using the mitochondrial gene matR (1,806 aligned base pairs). Overall, the estimated phylogenetic tree is highly congruent with independent analyses and provides a strongly supported placement of Rafflesia with the order Malpighiales, which includes poinsettias, violets, and passionflowers. Source: Jennifer Viegas, Discovery News, January 12, 2004; and various websites

http://radio.weblogs.com/0105910/2004/01/14.html

Mitochondrial DNA sequences reveal the photosynthetic relatives of Rafflesia, the world's largest flower

Proc Natl Acad Sci U S A. 2004 January 20; 101(3): 787–792. Published online 2004 January 8. doi: 10.1073/pnas.0305562101.

PMCID: PMC321759

Copyright © 2004, The National Academy of Sciences

Evolution

From the Cover

Mitochondrial DNA sequences reveal the photosynthetic relatives of Rafflesia, the world's largest flower

Todd J. Barkman,^*† Seok-Hong Lim,^* Kamarudin Mat Salleh,^‡ and Jamili Nais^§

^*Department of Biological Sciences, Western Michigan University, Kalamazoo, MI 49008; ^‡School of Environmental and Natural Resources Science, Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor, Malaysia; and ^§Sabah Parks, 88806 Kota Kinabalu, Sabah, Malaysia

^† To whom correspondence should be addressed. E-mail: tbarkman@wmich.edu.

Edited by Jeffrey D. Palmer, Indiana University, Bloomington, IN, and approved November 7, 2003

Received September 1, 2003.

Abstract

All parasites are thought to have evolved from free-living ancestors. However, the ancestral conditions facilitating the shift to parasitism are unclear, particularly in plants because the phylogenetic position of many parasites is unknown. This is especially true for Rafflesia, an endophytic holoparasite that produces the largest flowers in the world and has defied confident phylogenetic placement since its discovery >180 years ago. Here we present results of a phylogenetic analysis of 95 species of seed plants designed to infer the position of Rafflesia in an evolutionary context using the mitochondrial gene matR (1,806 aligned base pairs). Overall, the estimated phylogenetic tree is highly congruent with independent analyses and provides a strongly supported placement of Rafflesia with the order Malpighiales, which includes poinsettias, violets, and passionflowers. Furthermore, the phylogenetic placement of Mitrastema, another enigmatic, holoparasitic angiosperm with the order Ericales (which includes blueberries and persimmons), was obtained with these data. Although traditionally classified together, Rafflesia and Mitrastema are only distantly related, implying that their endoparasitic habits result from convergent evolution. Our results indicate that the previous significant difficulties associated with phylogenetic placement of holoparasitic plants may be overcome by using mitochondrial DNA so that a broader understanding of the origins and evolution of parasitism may emerge.

Methods Using the recent ordinal classification of angiosperms (11) as a guide, we explicitly sampled at least one family from 43 of 45 monophyletic orders to produce a broad mtDNA phylogenetic framework. Sampling included 92 species from 80 angiosperm families. Of primary interest for this study was the inclusion of Rafflesia keithii and its close parasitic relative Rhizanthes zippelii. Both of these species traditionally have been placed in Rafflesiaceae sensu stricto. Mitrastema yamamotoi (Mitrastemonaceae), another extreme holoparasite that has long been placed in Rafflesiaceae sensu lato, also was sampled to determine its phylogenetic affinities. Like Rafflesia, this parasite grows as an endophyte within its host (various members of the oak family, Fagaceae), emerging only during flower production (3). Because the morphology of Mitrastema is so divergent from that of photosynthetic plants, its affinities have remained obscure, with no relatives ever proposed aside from other endophytic parasites like Rafflesia (27). Three gymnosperms, Pinus, Ginkgo, and Zamia, were included as outgroups to root phylogenetic estimates. Table 1, which is published as supporting information on the PNAS web site, lists all of the species included in this study, the GenBank accession numbers for all of the sequences analyzed, and the voucher numbers for the newly generated sequences. Molecular methods, including DNA extraction, and DNA sequencing were performed as previously described (19). Details of the PCR methodology, including the primer sequences used, are available as Supporting Text, which is published as supporting information on the PNAS web site. In total, 55 matR mtDNA sequences were generated for this study. clustalx (28) was used to produce a preliminary alignment of the matR sequences that was followed by minor manual adjustments. The aligned data matrix is available from Tree-BASE (www.treebase.org/treebase) (S981–M1631). Regions of uncertain alignment were excluded before analysis; however, their inclusion did not alter the fundamental conclusions of this study. After exclusion, there were 1,499 included characters and 636 parsimony-informative characters. Unweighted parsimony analyses were conducted by using paup* v. 4.0b10 (29) with 100 random addition sequences, and tree bisection–reconnection swapping. Bootstrap support (BS) (30) and jackknife support (JS) (with 33% character deletion) values were obtained from 2,000 replicates by using “fast” stepwise addition. Although the fast stepwise addition analyses are expected to provide estimates of support that are less than those obtained when comprehensive branch-swapping analyses are performed (31, 32), such analyses were not computationally feasible with this data set. modeltest v. 3.06 (33) was used to determine the best-fit model of nucleotide substitution for the data set, and this best-fit model [K81 (34), assuming unequal nucleotide frequencies and a Γ parameter allowing for rate heterogeneity] then was implemented during Bayesian analyses performed by using mrbayes v. 3.0b4 (35). Four chains were run simultaneously for one million generations, and these were sampled every 100 generations. The first 10,000 generations were discarded as the “burn-in” period, and posterior probabilities (PP) for individual clades then were obtained from the remaining samples. Neighbor-joining (NJ) analyses were performed by assuming the optimal model of nucleotide substitution chosen with modeltest, and BS was obtained from 2,000 replicates by using paup* v. 4.0b10.

Results Parsimony analyses of matR from 95 species of seed plants resulted in 61,911 equal-length trees of 2,464 steps [consistency index = 0.5657]. The strict consensus tree (Fig. 1) represents an mtDNA-only estimate of broad, angiospermwide phylogeny, and it is largely congruent with estimates based on independent plastid and nuclear DNA sequences (9–11, 36). Levels of parsimony BS (P-BS) and Bayesian PP are shown on all nodes receiving estimates higher than 50 or 0.5, respectively. The complete majority-rule consensus tree obtained from the Bayesian analysis is available as Fig. 3, which is published as supporting information on the PNAS web site. Parsimony JS (P-JS) values and NJ-BS values could not be shown in Fig. 1 but are presented below. The complete set of JS values and the NJ tree with BS values are available in Figs. 4 and 5, respectively, which are published as supporting information on the PNAS web site. In general, the P-BS for all nodes obtained with fast stepwise addition was lower than the JS, NJ-BS, or PP and may be conservative. The P-BS values and PP shown for each node may therefore be interpreted as the lower and upper bounds of node reliability, respectively (37). The strict consensus tree suggests that the members of the ANITA (Amborella, Nymphaeales, and Austrobaileyales) clade (19, 22, 23) are the basal-most angiosperms, followed by an unresolved set of relationships among magnoliid orders, including all monocots. Expected major eudicot relationships include monophyletic rosids and asterids. All recently recognized orders are monophyletic when represented by more than one family, and, in most cases, BS and PP are high at the ordinal level (Fig. 1). Fig. 1. Strict consensus tree from an unweighted parsimony analysis of matR mtDNA sequences (number of trees = 61,911, tree length = 2,464, consistency index = 0.5657). BS from the parsimony analysis is listed before the slash, and PP from the Bayesian analysis (more ...) Within the context of this global angiosperm mtDNA phylogeny, the placement of Rafflesia as sister to the Malpighiales is supported (P-BS = 93, P-JS = 97, NJ-BS = 98, PP = 1.0). Parsimony analyses of amino acid sequences also suggested the same relationship (P-BS = 77, P-JS = 89) (Fig. 6, which is published as supporting information on the PNAS web site). The nested position of Rafflesia within the monophyletic Oxalidales + Malpighiales (P-BS < js =" 56," bs =" 57," pp =" 1.0)" href="http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=321759#ref10" class="cite-reflink">10, 11). Not supported by these data were traditional placements of Rafflesia + Rhizanthes with either Piperales (38) or Santalales (27). Placement of the enigmatic Southeast Asian holoparasitic plant Mitrastema (Mitrastemonaceae) with the Ericales (Fig. 1) was obtained by using the matR mtDNA phylogenetic framework produced here (P-BS = 70, P-JS = 82, NJ-BS = 91, PP = 1.0). This phylogenetic placement of Mitrastema with Ericales was surprising but is supported by parsimony analyses of amino acid data as well (P-BS = 76, P-JS = 91) (Fig. 6). Whereas Mitrastema has long been classified with Rafflesia + Rhizanthes, or close to them, these taxa are only distantly related.

Discussion

Holoparasite Phylogenetic Placement. The positions of the holoparasites, Rafflesia + Rhizanthes and Mitrastema, inferred with matR sequence data, are robust for several reasons. First, our results are not method-dependent, because the same strongly supported placements were obtained with parsimony, NJ, and Bayesian analysis. Second, long-branch attraction (39) does not seem to have misled these analyses, because the inclusion of random or misaligned sequences (40) did not influence the placement of Rafflesia + Rhizanthes or Mitrastema (Figs. 7 and 8, which are published as supporting information on the PNAS web site). Furthermore, it is clear that these lineages have not been artifactually placed with any of the other longest branches in this data set (Fig. 2A). Third, RNA editing and the potential for processed paralogy in plant mt genomes can confound phylogenetic estimates (41); however, it is unlikely that the placement of these holoparasites has been affected, because the removal of known matR RNA edit sites from the data set (42–44) did not change the level of support for the inferred relationships (Fig. 9, which is published as supporting information on the PNAS web site). Finally, because we have sampled the host plants of the parasites (or their close relatives), contamination cannot be invoked as an explanation for our results.

Fig. 2. (A) A randomly chosen phylogram of one of the equal-length trees from the matR parsimony analysis in Fig. 1. Branch lengths are shown only above lineages that had lengths of >50 steps. (B) Phylogram showing estimated 18S branch lengths on a topology (more ...)

Horizontal gene transfer (45, 46) of matR between Rafflesia and/or Rhizanthes and a member of the Malpighiales, or between Mitrastema and a member of the Ericales, is also an unlikely explanation for these results for several reasons. First, although host-to-parasite transfer of macromolecules is possible (47), Rafflesia and Rhizanthes are only parasitic on species of Tetrastigma (Vitaceae) to which they are clearly unrelated (Fig. 1). Likewise, Mitrastema only parasitizes members of the Fagales. Second, horizontal transfer seems to be rare in flowering plants overall (45) and would have to be postulated to have affected only Rafflesia, Rhizanthes, and Mitrastema, because all other nodes with high BS or JS in this analysis are comparably placed by independent estimates of angiosperm phylogeny (11). Finally, phylogenetic analysis of 128 coxI mtDNA sequences suggests comparable phylogenetic placements of Rafflesia with Malpighiales and Mitrastema with Ericales, although with lower support (C. W. dePamphilis, personal communication). Although confident placement of these parasites has been achieved with these data, more studies, including more taxa and characters, are needed to refine the positions of Rafflesia + Rhizanthes and Mitrastema to determine whether they are nested within, or sister to, the Malpighiales and Ericales, respectively. In either case, expanded circumscription of these orders is needed to include these holoparasites.

Rafflesia and Its Photosynthetic Relatives. The relationship between Rafflesia + Rhizanthes and Malpighiales estimated with matR mtDNA has not been suggested in any angiosperm classification or phylogenetic analysis. Not since 1822, when Robert Brown noted similarities in floral structure between Rafflesia and the Passifloraceae (12), has a potential relationship to the members of the currently circumscribed Malpighiales been suggested. Within the order, Rafflesia is morphologically most similar to Passifloraceae, with which it shares a hypanthium (perigone tube of Rafflesia), an androgynophore (central column of Rafflesia), and an annular corona (diaphragm of Rafflesia), a suite of characteristics that are otherwise somewhat rare in angiosperms (27). Some of these morphological features are found in other members of Malpighiales; therefore, the potential homology of these features may be ascertained only when the placement of Rafflesia, relative to a broader sampling of the order, is determined. The phylogenetic placement suggested by our data implies that a massive increase in flower size has occurred since the origin of Rafflesia, because the largest Malpighialian flowers are only 10 cm in diameter. Future comparative studies of MADS box and other floral developmental genes from Rafflesia and Malpighiales may reveal the genetic basis for this evolutionary floral size increase.

The placement of Rafflesia with Malpighiales also makes particularly tantalizing the 100-year-old observation that an individual of the tropical vine, Passiflora caerulea, was parasitic on another angiosperm (48). Although this may have been an aberrant observation of Passiflora, it is interesting to note that because Rafflesia exists as a vine-like endophyte within its host, the association of the viney growth form with parasitism is found in at least three separate orders: Laurales (49), Solanales (50), and Malpighiales. Although there may be different ancestral conditions giving rise to parasitism, the close physical association between vines and the plants that support them suggests that this interaction may foster the transition to a parasitic lifestyle.

The Asterid Holoparasite Mitrastema. The general placement of Mitrastema within the asterids has not been suggested previously, but its gamopetalous corolla supports this position. The specific placement of Mitrastema with the Ericales is further supported by a suite of morphological characteristics, including opposite and decussate leaves, parietal placentation, and circumscissile fruit dehiscence, that are also found in various members of the order but which collectively do not link it with any family in particular. The traditionally suggested close relationship between Rafflesia and Mitrastema is not supported by our data and probably reflects a taxonomic emphasis on their similar endoparasitic lifestyle. In fact, the floral morphology of Rafflesia is very different from Mitrastema, which has small (≈2.54 cm in diameter), white, bisexual flowers, a dehiscent staminal tube, and a superior ovary (Fig. 1). Because Mitrastema is no more divergent in floral morphology from Rafflesia than are any of the other traditional Rafflesiaceae genera, Cytinus, Bdallophyton, Apodanthes, and Pilostyles, it is possible that they represent independent parasitic lineages as well and should be targeted for future studies using mtDNA sequences.

Holoparasite mtDNA Evolution. The use of mt sequences to study parasitic plant phylogeny seems promising because the holoparasitic angiosperm family Hydnoraceae was recently placed by using a combination of mtDNA with plastid and nuclear sequences (51). One reason for the utility of mtDNA for studying the phylogeny of holoparasites is that the evolutionary dynamics governing these sequences seem to be similar in parasitic plants and their free-living relatives. Fig. 2 A shows reconstructed branch lengths from one of the equal-length trees obtained in the parsimony analysis of matR. It is clear that branch lengths are heterogeneous across angiosperms in general and that numerous photosynthetic plants are as divergent as Rafflesia or nearly so, yet all are confidently placed [except Ceratophyllum, which has defied confident placement to date (11)]. Furthermore, a likelihood ratio test (52) was performed to compare the level of selective constraint (estimated by the nonsynonymous/synonymous rate ratio, d_N/d_S) on matR in the holoparasitic lineages (Rafflesia, Rhizanthes, and Mitrastema) and photosynthetic lineages. Although the estimated d_N/d_S was higher for the holoparasitic lineages (1.0646) than for the photosynthetic lineages (0.7010), this difference was not statistically significant (P > 0.05). Thus, although some holoparasitic plants show elevated substitution rates and significant decreases in selective constraint for chloroplast genes such as rbcL and rps2 compared with their photosynthetic relatives (5, 6), matR has likely been maintained for efficient mt functioning even in parasitic plants.

Problems with Holoparasite 18S Nuclear DNA. In addition to mtDNA, nuclear sequence data could provide an important independent genomic estimate of holoparasitic relationships as well. However, the 18S nuclear ribosomal DNA and other loci involved in protein translation have been problematic in studying other extreme parasites like microsporidia (2), and these genes are unlikely to provide useful data for Rafflesia or most other plant holoparasites. To illustrate this point for Rafflesia (Mitrastema has not yet been sequenced), we have obtained 70 nuclear 18S DNA sequences from GenBank for taxa that are also represented in our matR data set. Table 2, which is published as supporting information on the PNAS web site, lists all species names and GenBank accession numbers. Fig. 2B shows branch lengths for the 18S sequence data on a tree that was constrained to represent the currently accepted estimate of angiosperm relationships (10, 11). Assuming a placement of Rafflesia as sister to the Malpighiales, it is clear that this holoparasite has a highly divergent 18S sequence that is 4 times longer (197 steps) than the next longest sequence in the data set (the branch separating angiosperms and gymnosperms; 49 steps), and it is 6–8 times longer than most other branches. As expected with long-branch attraction, Rafflesia is placed with the other longest branch in the data set in an unconstrained parsimony analysis (Fig. 2C). This result corroborates an earlier study of 18S that found Rafflesia artifactually placed as sister to angiosperms and indicates that this gene is of limited utility for studying this divergent holoparasite (18). Even phylogenetic analyses of the slowly evolving 16S plastid ribosomal DNA clearly show extreme divergences in holoparasites (4), indicating that this gene will also be of limited utility in studying parasitic plants like Rafflesia. Classes of nuclear gene sequences other than those involved in translation, such as loci involved in primary metabolism, may instead provide better candidates for studying holoparasite relationships (2). Future studies of nuclear genes from Rafflesia and other extreme holoparasitic plants are needed because no other loci besides the 18S ribosomal DNA have been sequenced.

Implications. The results of this study have clear implications for studies of angiosperm phylogeny and parasite evolution. First, we show that mtDNA sequences provide a third independent genomic estimate of angiosperm relationships useful for resolving a variety of phylogenetic questions among both anciently and more recently evolving lineages. Second, placement of several other parasitic plants [composing nearly half of the taxa that remain of uncertain positions at the ordinal level (11)] may now be possible using the approach adopted here. Third, the eventual phylogenetic placement of all parasitic plants within flowering plant phylogeny will enable detailed comparative studies aimed at inferring ancestral states that may have facilitated the repeated evolution of parasitism in angiosperms.

Supplementary Material

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Notes

This paper was submitted directly (Track II) to the PNAS office.

Abbreviations: mt, mitochondrial; BS, bootstrap support; JS, jackknife support; PP, posterior probabilities; NJ, neighbor joining; P-BS, parsimony BS; P-JS, parsimony JS.

Data deposition: The sequences reported in this article have been deposited in the GenBank database (accession nos. AY453070–AY453124).

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