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Contents
Classification of Phalaenopsis
Pollinia
Amabilis Complex
Variant Species
Cornu-cervi Complex
Lueddemanniana Complex
P. fasciata/P. reichenbachiana
P. lueddemanniana/P. delicata
P. violacea/P. mentawai
P. sumatrana Complex
P. gigantea Complex
P. equestris
 Under construction for no-linked items

Classification and Specificities of Genus Phalaenopsis

 In recent years, analysis of molecular data has been used to classify the genus Phalaenopsis. Furthermore, analyses of the chloroplasts and nuclear ribosomal DNA have clarified the phylogenic and evolutionary trends. The latest classification, however, still leaves some inconsistencies. This might stem from investigations done with a limited number and regionality of samples in the DNA analysis. The latest classification of genus Phalaenopsis was proposed by E.A.Christenson and comprises 62 species, dividing the genus into five subgenera: - Proboscidioides, Aphyllae, Parishianae, Polychilos, Phalaenopsis, - and subdividing two subgenera of Polychilos and Phalaenopsis into four sections.

 Although further studies in scholarly classification must be relied by botanist, orchid-collectors have often come across imported plants species that are hard to identify due to, e.g., similar appearance (lueddemanniana and cornu-cervi complexes), regionally different morphologies (amabilis complex), and hybrid-like varieties. Sometimes, unknown native species are found in the catalog of southeast asian nurseries. Even from a professional point of view, classification is still one of the unsolved issues in some groups of native species. This page posts a few Phalaenopsis species that have taxed grower's brains with this sort of problem.

 The visual information for identifying Phalaenopsis species are as follows:

  1. perianth shape and color
  2. callus structure
  3. post-pollination chlorophyll of perianth
  4. scent
  5. leaf morphology (shape, color, and texture)
  6. inflorescences (shape, color, and length of stalk)

 Due to strong individual characteristics in the native species, the difference between most species can be easily recognized by looking at the perianth shape and color. However, when collecting similar species, e.g., lueddemanniana complex, amabilis complex, sumatrana complex, or fuscatae complex, and their varieties or sister species, hobbyists may have difficulty in identifying them from the appearance of the perianth due to the complexities in individual and area varieties. Furthermore, the hobbyists may only infrequently find characteristics that are different (color, shape, scent of flower) from others even within the same lot or may come across mislabeled species. To cope with such problems, it is necessary to have some knowledge of the callus structure in the lip and the differences in morphological features between sister species.

 Genus Phalaenopsis is classified into two groups that have four- and two-pollinium clades. The species with four pollinia in the subgenera Proboscidioides, Aphyllae, and Parishianae are distributed in south China, India, northern Vietnam, Myanmar and Thailand, respectively. The subgenera Polychilos and Phalaenopsis are mainly distributed in Indonesia, Malaysia and the Philippines, as shown in Figure 1. P. deliciosa with four pollinia and P. cornu-cervi, P. sumatrana, and P. kunstleri with two pollinia are distributed in both areas.


Fig. 1 Distribution Map of Two- and Four-Pollinia Groups

 Over the last decade, the evolutionary phylogenic tree and the genetic distance of genus Phalaenopsis have been inferred by DNA analysis, and the relationships among the species are also being clarified. However, some hobbyists who have acquired various species including varieties and inexplicably similar species must be feeling that the DNA samples must be inadequate (due to, e.g., individual specificity, areal difference, variety, differences in scent, etc., in identical species).


Varieties with Two and Four Pollinia

 Until recently, many species of subgenera Aphyllae (wilsonii, braceana, minus, hainanensis, etc.), Proboscidioies (lowii), Parishianae (appendiculata, gibbosa, lobbii, parishii) and section Deliciosae (chibae, deliciosa, mysorensis) had been classified in the genus Kingidium, which has distinctive morphologies - four pollinia, small flowers, deciduous or the largest lateral sepals in the tepals - from other subgenera. It was the late 1990s that they were included in the genus Phalaenopsis. According to the phylogenic tree of the genus, subgenus Aphyllae is regarded as a root base and has developed to Proboscidioies, Parishianae and section Deliciosae, in that order. Based on this presumption, one theory states that subgenus Aphyllae is the oldest ancestor of genus Phalaenopsis and that it moved southward from the Himalayas to Indochina, Malaysia, Indonesia and the Philippines.

 Figure 2 shows the pollinia of P. deliciosa and P. chibae in section Deliciosae (left two images), P. amabilis in section Phalaenopsis (second from right), and P. violacea (far right) in section Amboinenses. The former two sections are grouped in the subgenus Phalaenopsis, while P. violacea belongs to the subgenus Polychilos. Four flattened pollinia can be observed in the left two, and two rounded and thick pollinia in the right two, respectively.


P. deliciosa         P. chibae

P. amabilis           P. violacea
Fig. 2 Four and Two Pollinia

 Figure 3 shows P. wilsonii, P. hainanensis, and P. minus in the subgenus Aphyllae. In the pictures, the perianth of P. minus has a distinctive appearance. It is hard to understand the theory why P. minus is classified in the same subgenus Aphyllae comprised of P. wilsonii, and P. hainanensis despite having a very different shape from the other species. DNA analysis indicates that P. minus is somewhat genetically distant from the others and might be older (more primitive) in the phylogenic tree.


P. wilsonii

P. hainanensis

P. minus
Fig. 3 Species in Subgenus Aphyllae

 The species with the most unique shape of perianth are those in the subgenus Parishianae, as shown in Figure 4. The lateral sepals are prominently larger and wider than the dorsal sepal and petals. The lip has a triangular shape, and a mobile midlobe sways back and forth, which is the only characteristic that specifies it as being in the genus Phalaenopsis. The visual impression of the midlobe including its movement seems to be mimetic of an insect opening its wings, and the filiform callus looks like the antenna. When the insect overrides the midlobe and takes off, the head of the insect strikes against the rostellum of the column, and the pollinia is removed by sticking to the insect's head. At the next stage involving the same behavior, the pollinia on the head are caught by the talon of the column, and the pollination is completed. This is just speculation, however, based on the structure of the midlobe, although it may be worth considering. It would be interesting to see what kinds of insects visit these flowers.


P. appendiculata

P. parishii

P. gibbosa
Fig. 4 Subgenus Parishianae 

 Another inexplicable matter in the latest classification is that the section DeliciosaeP. chibae, deliciosa, mysorensis) and the section Phalaenopsisamabilis, aphrodite, schilleriana, stuartiana, etc.) are grouped in the same subgenus Phalaenopsis, despite the fact that the two have remarkably different morphologies, as listed in Table 1.

Table 1 Morphologies of Sections Deliciosae and Phalaenopsis
 
Deliciosae
Phalaenopsis
Largest sepals and petals
lateral sepal
petal
Sizes of sepals and petals
1.2-1.3cm
5-9cm
Pollinia
four
two
Callus
uniseriate (bifit) or biseriate
uniseriate (shield-shape)
Leaf

thin, dark green, undulation

thick, cuticular wax
Lip midlobe
as shown in Figure 4
triangular
Midlobe apex
smoothed convex
flexuous tendril

 According to the DNA analysis, the section Deliciosae has some genetic distance from the sections Phalaenopsis and Polychilos (amboinensis, gigantea, lueddemanniana, violacea, etc.). This might imply that the perianth of species in the section Deliciosae has a remarkably different morphology from those in the section Phalaenopsis. In contrast, P. chibae and P. deliciosa have a similar appearance to the species in the section Parishianae and section Aphyllae, respectively, as shown in Figs. 3 and 4. Because of this, some botanists have proposed that that the section Deliciosae be classified into the subgenus Aphyllae. Considering the four pollinia structure of section Deliciosae, it seems to be a convincing proposal. One of the reasons why the section Deliciosae and Phalaenopsis should be grouped into the same subgenus Phalaenopsis is said to be as follows: "The species in Deliciosae are not deciduous and do not exhibit post-pollination chlorophyll of the perianth, which differ from the species in Aphyllae." However, the species in Aphyllae also have prognostic characteristics such as not defoliating in warmer climates throughout the year. Furthermore, it is not correct that the species of Aphyllae exhibit post-pollination chlorophyll of the perianth; rather, the perianth withers after pollination (see the P. wilsonii or P. honghenensis in the page on "58 species"). Thus, the above contention is questionable.


P. chibae

P. deliciosa

P. amabilis

P. schilleriana
Fig. 5 Sections Deliciosae and Phlaenopsis

 In the book by C. Singchi, T. Zhanhuo, et al. called Native Orchids of China in Color 2d ed. Science Press, 2006), the authors state that P. mannii and P. wilsonii are classified in the genus Phalaenopsis but P. braceana is still grouped into Kingidium. In the Philippines, P. deliciosa is called Kingidium Philippinense in the market.

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P. Amabilis Complex 

 The P. amabilis group is widely distributed throughout Taiwan, the Philippines, Borneo, Indonesia, Malaysia, the Maluku Islands, and Australia. The P. amabilis group comprises P. amabilis, P. aphrodite, P. philippinense, P. sanderiana, P. schilleriana, and P. Stuartiana, which have common features of relatively large sepals and petals, a long flower stalk, thick leaves and roots, and they do not exhibit post-pollination chlorophyll of the perianth. P. amabilis and P. aphrodite, in particular, have deep similarities and are hard to identify from their appearance. Only the calluses have different features. The species of the amabilis group, except for P. amabilis, are endemic to their habitats in the Philippines, as listed in Table 2.

Table 2 Habitats of Amabilis Group
P. amabilis
Palawan, Twai Twai island
P. aphrodite
Bataan, Camiguin, Luzon, Mindanao, Bilirau, Leyte, Samar, Negtros, Babuyan Is.
P. philippinense
North Philippines (Luzon)
P. sanderiana
South Philippines (Mindanao, Igat, Balut and Saragani)
P. schilleriana
Luzon
P. stuartiana
Mindanao, Leyte

 

 According to the genetic affiliation, the amabilis group is further divided into two subgroups, amabilis and schilleriana complexes. The amabilis complex comprises P. amabilis, P. aphrodite and P. sanderiana, while the schilleriana complex includes P. philippinense, P. schilleriana and P. stuartiana. The oldest ancestor (root of the evolutionary phylogenic tree) is inferred to be P. aphrodite, which is widely distributed in the Philippines. Other species have more limited habitats. Some of them make their habitats together but have different blooming seasons.

 There is one theory about the evolutionary process and distribution of the amabilis complex in Southeast Asia, as follows: first, P. aphrodite was brought into the region at Palawan and evolved into P. amabilis. Thereafter, P. amabilis dispersed into Borneo, Sumatra and Malaysia. In contrast, P. aphrodite dispersed to Taiwan through the Calayan islands and the southern Philippines. It evolved into P. sanderiana in southern Mindanao from P. amabilis. P. amabilis further dispersed into Sulawesi and New Guinea, developing into P. amabilis moluccana and rosenstromii respectively. Thus, P. amabilis in Borneo and Sulawesi came along different pathways: the respective Palawan and Mindanao routes.

 One question arises in the above theory about whether it is stochastically possible for P. amabilis to evolve from different species (P. aphrodite or P. sanderiana) through different places (from Palawan to Borneo, and from Mindanao to Sulawesi). In the route from Palawan to Borneo, the species exhibiting an intermediate feature (form) between P. amabilis and P. aphrodite can be observed in Palawan. In contrast, P. amabilis and P. amabilis-like species cannot be found in either Luzon or the Mindanao islands. Furthermore, there is no P. aphrodite nor P. sanderiana in Sulawesi. P. aphrodite in the southern Philippines and P. sanderiana in Mindanao have a similar callus form, but they are not similar to P. amabilis, which indicates that thre is no evidence that P. aphrodite evolved into P. amabilis in Luzon or the Mindanao islands. Reasonable evidence might be necessary to believe the assumption of an evolutionary process through the Mindanao route.

 P. sanderiana is reported to be genetically closer to P. amabilis than to P. aphrodite. This is also problematic because the callus structure of P. sanderiana is very similar to the callus of P. aphrodite in the southern Philippines, and is not very similar to P. amabilis, as described above.

 Although not based on a scholarly background, it is nonetheless interesting to imagine the evolutionary process of plants from a morphological point of view, just like unraveling the mystery of an ancient monument. Therefore, I state one assumption on such a standpoint as follows:

  1. First, P. aphrodite, which was distributed around Palawan, evolved into P. amabilis. Thereafter, the territory of P. amabilis grew to include Sabah Borneo, while P. aphrodite in Palawan moved toward the north and dispersed into Taiwan through the Calayan islands. P. aphrodite in the Philippines moved further southward and spread throughout the Philippines.
  2. P. amabilis in Sabah Borneo developed various forms, e.g., a long midlobe, or dark green or partly brownish-red leaves as seen in the form of grandiflora; thereafter, it dispersed to three regions - Sumatra, the Sulu-Archipelago and Sulawesi.
  3. P. amabilis in Sumatra dispersed into Malaysia and the Mentawai Islands. P. amabilis in Sulawesi dispersed into Java, East Timor, Maluku Islands and New Guinea, while P. amabilis in Sulu-Archipelago expanded into southern Mindanao and evolved into P. sanderiana

 The reason P. amabilis cannot be found in most Philippine islands except for Palawan and the island at the tip of the Sulu-Archipelago, despite its wide distribution in Southeast Asia, is assumed to be as follows: Palawan, Mindanao and Zamboanga are old islands in the Philippines that were once connected to Borneo on the same Eurasian continent before 10Mya. On the contrary, most of the other Philippine islands are young and came along after 5Mya. The emergence of P. amabilis that evolved from P. aphrodite in Palawan occurred at the same stage of crustal movement separating these islands from the continent. Therefore, P. amabilis could not move across the inland Philippines.

 On the other hand, the Malay Peninsula, Borneo, Sumatra, Java, and western Sulawesi on the Sunda Shelf would have been interconnected during glacial times at a lowered sea-level, and this would have enabled P. amabilis to disperse into these regions. P. sanderiana has a restricted distribution in southern Mindanao, and it is said to be genetically closer to P. amabilis than to P. aphrodite. P. amabilis does not inhabit southern Mindanao. This can be explained by the fact that the P. amabilis from Sabah moved to the Sulu-Archipelago at the age when Borneo and Zamboanga were connected, and it evolved into P. sanderiana in the eastern part of the Sulu-Archipelago. After Zamboanga separated from Borneo and the Sulu-Archipelago, P. sanderiana remained in parts of southern Zamboanga and Mindanao. Thus, P. amabilis is found in the southwestern tip of Sulu-Archipelago, and P. sanderiana is found in the islands of Zamboanga, but not found P. amabilis in Mindanao.

 Looking at the Google map (satellite) around the Philippines and Borneo Island, Palawan and the Sulu-Archipelago islands lay in-between Borneo, and the Phiippines are in the shallow sea. On the otherhand, there is a deep trench between Borneo and the Sulawesi Islands. This might be the main reason supporting the theory of a Mindanao route (from Glan, Mindanao to Manado, Sulawesi). However, the distance between the shallow sea off Kalimantan Selatan and Manuju, Sulawesi is 20 km, and there is a sea studded with small islands between Kalimantan Selatan, and Makassar, Sulawesi.

 The above hypothesis originated from one scholarly theory that P. amabilis evolved from P. aphrodite. However, if this process occurred in the reverse order, some inconsistencies in that scholarly theory and my theories could be demystified by the following scenarios; First P. amabilis appeared in Borneo, dispersed into Sumatra, Sulawesi, Sulu-Archipelage and Palawan, evolved into P. aphrodite in Palawan, and evolved into P. sanderiana in the eastern part of the Sulu-Archipelago. P. aphrodite further dispersed into the Philippines and Taiwan, while P. amabilis dispersed into Malaysia, Java, and Australia. If we consider the current distributions of the P. amabilis group in SE Asia and the history of continental drift, this theory looks more compelling than the previous one.


P. amabilis

P. aphrodite

P. philippinensis

P. sanderiana

P. schilleriana

P. stuartiana
Fig. 6 P. amabilis Group

 As the result of out-migration and evolution, P. amabilis developed into various subspecies: moluccana (Sulawesi) and rosenstromii (New Guinea). P. amabilis subsp. rosenstromii has a certain genetic distance from P. amabilis in other areas. DNA analysis indicates that the genetic distance between P. aphrodite in Palawan and Borneo is very short. This can be estimated due to the geographical relationship of these areas. Furthermore, intermediate forms can be observed on the lips of P. aphrodite and P. amabilis.


P. amabilis Palawan

P. amabilis Borneo

P. amabilis Borneo

P. amabilis Sumatra

P. amabilis Java

P. amabilis Irija
Fig. 7 P. amabilis in Palawan、Borneo、Sumatra, Java、and Irian Java

P. amabilis Palawan

P. amabilis Borneo

P. amabilis Borneo

P. amabilis Sumatra

P. amabilis Java

P. amabilis Irija
Fig. 8 Lip Structures of P. amabilis (removed lateral lobes)

  Due to the above evolutionary backgrounds, Phalaenopsis species in Palawan and the Sulu-Archipelago, which were historically connected to Borneo, are very important regions in which to study the evolutionaryl morphologies of the species. However, these species are unobtainable now because of regulations against extracting plants from the wild.

Fig. 9 Lip and Callus Structures of P. aphrodite (removed lateral lobes)

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Variant Species in P. amabilis Complex

1. P. amabilis ? from the southern part of the Philippines:

 Figure 11 shows one of three species imported from the Philippines under the pretext of being "P. amabilis." It is apparently different from P. amabilis (mislabel) but similar to P. aphrodite from the view of the callus structure. The leaf morphologies are quite similar to P. amabilis. Most P. aphrodite plants, as seen in Figure 9, have a callus that is rather rounded and thick; however, this callus has a sharp edge and is thin, which is similar to the callus of P. sanderiana. The habitat is not clear, but it is estimated to be in the southern part of the Philippines.

Fig. 11 P. aphrodite Imported on Pretext of being P. amabilis

2. P. amabilis from Palawan:

 To specify the unique features of P. amabilis from Palawan, this section addresses the differences in their morphology. Figure12 (A - E) shows the callus shapes of 10 species named P. amabilis Palawan, while two species in Figure 12 F are well known and typical P. amabilis species from other places.


A

B

C

D

E

F
Fig. 12 P. amabilis (A-E) from Palawan and P. amabilis (F) from Borneo

 The 10 species shown in A - E were shipped in the same lot under the label P. amabilis Palawan, so they are assumed to be from the same habitat. From the pictures, it looks as if many of the species are not P. amabilis but P. aphrodite. The callus shapes can be classified into three types - similar to P. amabilis at A(L), B(R), and D(R), an intermediate form of P. amabilis and P. aphrodite at B(L) and D(L), and similar to P. aphrodite at C and E - , in which (L) and (R) denote left and right positions in each picture. Figure 13 shows the top view of lips in A - E (left picture), and the difference between P. amabilis Palawan (upper row in right picture) and a typical P. amabilis (lower row in right picture)

Fig. 13 Callus and Midlobe Structures

 Figure 14 shows close-up images. The calluses from left to right in the top row are E(L), A(L), and D(R), which are very similar to the forms of P. aphrodite, P. amabilis and intermediary body between them, respectively. Pictures (J) and (K) are typical P. amabilis from Java. Picture (L) is the side of the callus in picture (I).


G

H

I

J

K

L
Fig. 14 Palawan amabilis (G、H、I、L) and P. amabilis (J、K)

 Figure 15 shows typical callus shapes of P. aphrodite. As shown in the picture (far left), the callus has shield-shaped, notched edges at the right and left with three convex curves (denoted as 1, 2 and 3) on each side. The posterior part looks like two carina plates (A and B in far right picture) conjugated at the anterior edge. In contrast, P. amabilis does not have such a structure, and further, the convex curve denoted by 3 (far left picture) is very small or not observed at all.

Fig. 15 P. aphrodite Calluses

 From that view point, again, observing the callus shapes in Fig. 14, G, H and I are similar in shape to P. aphrodite, P. amabilis and intermediary body, respectively. H is similar to the calluses of J and K. K is P. amabilis v. grandiflora (Borneo). Figure 16 shows P. aphrodite、P. amabilis Palawan、P. amabilis Palawan and P. amabilis grandiflora from left to right.


Fig. 16 P. aphrodite, P. amabilis Palawan, P. amabilis Palawan, P. amabilis grandiflora (left to right)

 Figures 12 - 16 show the various callus shapes of P. amabilis from Palawan, which includes a mixture of P. aphrodite- and P. amabilis-like species. The morphological variation as seen above within the narrow area may be peculiar in Palawan or the Sulu Archipelago, where the evolutionary species grew before 10Mya.

3. P. amabilis Clump type:

 Figure 17 shows P. amabilis v. formosana imported under the pretext of being "clump" type plants. This plant develops many leaf stalks from a main culm, like a sympodial plant, which overturns the monopodial theory of genus Phalaenopsis. The lateral bud of flower stalk may mutate into a leaf stalk due to any cause. This characteristic is also observed in P. cornu-cervi and P. x intermedia. According to the latest classification, it is said that P. amabilis does not inhabit Taiwan and previously identified P. amabilis v. formosana should be grouped into P. aphrodite as subsp. formosana. Observing the callus structure, it doesn't have a definitive shape like P. amabilis nor P. aphrodite. The lip shape of the fan-like form is also unique.

Fig. 17 P. amabilis v. formosana Clump Type

4. Phal. amphitrite ? :

 Figure 18 shows P. stuartiana with pale pink dorsal sepal and petals, which differs from the original white color. It appeared in only one of 300 plants shipped from Mindanao island. Observing the color shade, it looks a natural hybrid of P. stuartiana and P. schilleriana. However, P. schilleriana does not inhabit Mindanao. Another possibility is that it is a hybrid with P. sanderiana, although there is less chance for a cross due to the different blooming season. Other morphologies, e.g., leaves and flower stalk, are quite the same as P. stuartiana. If this is a hybrid of P. stuartiana x P. sanderiana, it is a P. amphitrite and may be the world's first color photograph of a natural hybrid since its discovery in 1900.

Fig. 18 P. stuartiana SP

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P. Cornu-cervi Complex

 The Cornu-cervi complex is classified in the subgenus Polychilos comprising P. borneensis, P. cornu-cervi, P. lamelligera, P. mannii, and P. pantherina, as shown in Figure 19. Many people are of the opinion that P. lamelligera does not have a definite base to be identified as a different species from P. cornu-cervi. However, P. lamelligera is a genetically distant from P. cornu-cervi, which has been verified by recent DNA analysis. Although P. cornu-cervi is a more widely distributed than P. amabilis, it is not as well-known as P. amabilis, which defines the subgenus and regional varieties.

Table 3 Habitat of Cornu-cervi Complex
P. borneensis
Borneo
P. cornu-cervi
Northeastern India, Thailand, Nicobar Is, Bhutan, Java, Borneo, Sumatra, Philippines
P. lamelligera
Borneo
P. mannii
Northeastern India, Nepal, China, Vietnam
P. pantherina
Borneo


P. borneensis

P. cornu-cervi

P. lamelligera

P. mannii

P. pantherina
Fig. 19 P. cornu-cervi complex and Callus Shapes

Thailand

Java

Philippines
Fig. 20 Regional Differences of P. cornu-cervi

 From a morphological point of view, P. cornu-cervi has many varieties of lip structure, perianth, scent and stem, but surprisingly few enthusiasts have discussed this species. It can be divided into four forms: Thailand, Borneo, Java and the Philippines. The apex shape of the midlobe varies remarkably depending on the habitat as shown in Fig. 20. P. cornu-cervi in Thailand, Sumatra, and Borneo continue blooming for around half a year; however, P. cornu-cervi from the Philippines blooms one by one all year round in the greenhouse (strictly speaking, there is a rest period of a few months for flowering in early spring), which is the only species (except P. equestris in Mindanao) to do so in genus Phalaenopsis. It is often observed to grow in clusters in the Philippines, and to have a scent in Java. Other plants with clustered growth are also seen in P. amabilis, P. x intermidia and P. lueddemanniana.

 It remains to be clarified which species is the ancestor of the P. cornu-cervi complex. Looking at the evolutionary phylogenic tree of section polychilos, P. mannii is considered to be the oldest species, and the next is P. lamelligera. P. cornu-cervi is the third. P. mannii is distributed throughout a wide range from India to Indochina, just as P. cornu-cervi is. If P. mannii was the ancestor of P. cornu-cervi, it might have come from around the Himalayas and evolved into P. cornu-cervi. Thereafter, P. cornu-cervi dispersed into Indochina, Malaysia, Borneo and the Philippines. The P. cornu-cervi on Borneo island subsequently evolved further into P. borneensis and P. pantherina. Both P. borneensis and P. pantherina inhabit Borneo, which suggests that these are new species that evolved after separating from Borneo island from the Eurasian continent during the Pleistocene epoch.

 There is a question about the classification in the section Polychilos in regards to why P. mannii, with its cylindrical flower stalk, is grouped into the same section in which all other species have a flattened zigzag form, as shown in Figure 21. The same problem is found with P. violacea (Sumatra and Mentawai). The second question is why P. lamelligera of the second generation inhabits only Borneo even though P. cornu-cervi of third generation is distributed in almost the same region as P. mannii, which shows the contradiction of the evolutionary process as P. mannii -> P. lamelligera -> P. corn-cervi. The third question is about the scent. P. cornu-cervi is noted as a non-scented species. However, P. cornu-cervi plants imported from Indonesia are often found to have a strong scent. The scent is a very important factor to attract pollinators. Whether a plant has a scent or not must be strongly associated with the historical development and environmental background of the species. This characteristic of the same species sometimes having a scent and sometimes not having one is also found in P. modesta and P. hieroglyphica. However, threre has been very little investigation on scent in the taxonomy of genus Phalaenopsis. There is a possibility that it may be a new species that has not been classified yet.


P. cornu-cervi

P. mannii
Fig. 21 Flower Stems of P. cornu-cervi andP. mannii

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Lueddemanniana Complex

 Lueddemanniana complex is classified to the section Amboinenses and comprises eight species - P. bastianii, P. fasciata, P. hieroglyphica, P. lueddemanniana, P. mariae, P. pallens, P. pulchra, P. reichenbachiana - which inhabit the Philippines except P. mariae (Borneo and the Philippines). Table 4 lists the habitat of the Lueddemanniana complex in the Philippines.

Table 4 Habitat of Lueddemanniana Complex
P. bastianii
Sulu Archipelago
P. fasciata
Luzon, Mindanao, Bohol
P. hieroglyphica
Palawan, Polillo, Mindanao
P. lueddemanniana
Appari, Mindanao
P. mariae
Luzon, Mindoro, Mindanao, Sulu-Archipelago, (Borne Is)
P. pallens
Luzon, Province Bataan, Mt Mariveles, Lanao River
P. pulchra
Luzon, Leyte
P. reichenbachiana
Mindanao

P. bastianii

P. fasciata

P. hieroglyphica

P. lueddemanniana

P. pallens

P. pulchra

P. mariae

P. reichenbachiana
Fig. 22 Lueddemanniana Complex

 The Lueddemanniana complex except P. mariae has a restricted habitat only in the Philippines. Geological records indicate that Borneo, Palawan and Zamboanga were on the same Eurasian Plate 10Mya, and Palawan and Zamboanga separeted from Borneo about 5Mya. Against this background, P. mariae found in both Borneo (Kalimantan) and the Philippines is assumed to be the oldest species among lueddemanniana complex. P. mariae has been investigated as the origin of the lueddemanniana complex through studies of the evolutionary phylogenic tree. It is interesting to find P. mariae in the Sulu Archipelago (Jojo island) and Borneo Kalimantan. This might indicate the movement of P. mariae from Borneo to Mindanao and vice versa through Sulu Archipelago. Thus, P. amabilis and P. mariae moved between the Philippines and Borneo at the same era through Palawan and Sulu Archipelago respectively. It is assumed that seven other species are new species that have evolved from P. mariae during the Pleistocene epoch (5 Mya).

 In the Lueddemanniana complex, the genetic distance is closer between P. bastianii and P. Pallens, between P. hieroglyphica and P. reichenbachiana, P. fasciata and P. pulchra. However, the callus structures are analogous between P. hieroglyphica and P. fasciata, and P. reichenbachiana and P. pulchra. Some taxonomist have suggested that P. reichenbachiana may possibly be a natural hybrid between P. fasciata and P. hieroglyphica due to the certain genetic distance from the others. P. lueddemanniana v. delicata is well known as the variety of P. lueddemanniana. However, both have a unique morphology, as described below.

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P. fasciata and P. reichenbachiana

 P. fasciata and P. reichenbachiana are species that are extremely difficult to identify only from the appearance, which is similar to P. fuscata and P. kunstleri. P. fasciata is widely distributed from Luzon to Mindanao islands, while P. reichenbachiana is limited to Mindanao. To my knowledge, there is no Philippine nursery that consciously grows these two species separately; they usually cultivate them together as P. fasciata.

 According to the illustration by H. R. Sweet shown in Figure 23, P. fasciata has a midlobe that is oblong-obovate, an apical raised pad with a few scattered trichomes, the callus biseriate, the anterior callus bifid and the posterior callus with densely tubercles, while P. reichenbachiana has a midlobe that is trowel-shaped with acute, lateral margins that are irregularly serrulate, the callus triseriate, the anterior and central callus bifid, and the posterior callus tubercles. Although P. reichenbachiana has sepals and petals that are more rounded and ovate than P. fasciata, the perianth shape can not be the definitive factor to identify the species, because both may have similar shapeas with individual variations. After observing more than 50 samples of P. fasciata from the Luzon and Mindanao regions, I found that Reichenbachiana-like species with the above features are often included from the Mindanao region. However, the problem is that various (intermediate) morphologies are observed in both regions.


P. fasciata Callus

P. Reichenbachiana Callus
Fig. 23 Lip Structure of P. fasciata and P. reichenbachiana (Illustrated by H.R. Sweet)

 The remarkable difference in Fig. 23 is the midlobe structure. However, common shapes are observed in various species; not only P. fasciata but also, e.g., P. hieroglyphica and P. lueddemanniana. Therefore, this feature is difficult to use as the definitive factor for identification. Up to the present, three distinctions have been noted in P. reichenbachiana.

  1. rounded sepals and petals
  2. midlobe trowel-shaped
  3. central callus with bifid

Furthermore, E.A. Christenson pointed out that P. fasciata has a green apple fragrance, while P. reichenbachiana has a musty odor. As described above, a single determinant is not enough to identify them. It may need multiple coincides for three determinants above. Fig. 24 shows two species similar to P. fasciata and P. reichenbachiana.


P. fasciata

P. fasciata

P. fasciata

P. reichenbachiana

P. reichenbachiana

P. reichenbachiana
Fig. 24 Perianth and Callus Structures of P. fasciata and P. reichenbachiana

  Figure 25 shows two species imported from Luzon island with the name of P. fasciata. In the picture, the species (far left) has sepals and petalsthat are oblong-lanceolate, while the species (third right) has sepals and petals that are oblong-obovate, which is similar to P. reichenbachiana. However, both species came from Luzon, which is not a habitat for P. reichenbachiana. On the other hand, the midlobes of both species show oblong-obovate and apical raised pads without trichomes, which is a characteristic of P. fasciata. In the callus structure, both have bifid anterior and tubercles on the posterior callus. However, small bifid denticulation at the midportion between the anterior and poster callus is observed in the left species. In contrast, the right species has a clearly recognizable and protruded bifid denticulation at the midport (far right). The left species may include an apophysis at the midport to the posterior callus. It is not certain if the right species does. These morphologies might represent the intermediate feature between P. fasciata and P. reichenbachiana.

 Meanwhile, many P. fasciata imported from Mindanao have sepals and petals that are oblong-obovate as shown at the right (third right) of Fig. 25 and that have two types of midlobe shape, as shown in the lower far right picture of Fig. 24 and in the far right picture of Fig. 25. Furthermore, it does not emit a green apple scent but a slightly fusty smell.

P. fasciata
P.fasciata (imtermediate type)
Fig. 25 P. fasciata (left) and Intermediate Species (right) of reichenbachiana and fasciata

 Figure 26 shows a P. reichenbachiana-like species from Mindanao island. The sepals and petals are obovate, and the midlobe is trowel-shaped, similar to the P. reichenbachiana illustrated by H. R. Sweet. The bifid denticulation at the midport is identified as a callus that is independent from the anterior and posterior callus, and therefore, this species is P. reichenbachiana with a triseriate callus. Looking at the far right picture, the central boss seems independent from the other two.

 P. fasciata imported from Mindanao (2008 - 2010) have never been observed with the definite features of P. fasciata shown in the top row of Fig. 17, but rather with rounded sepals and petals and an intermediate structure of midlobe and callus. On the other hand, P. fasciata from Luzon have some rounded sepals and petals, but most have the midlobe shape shown in the second left image of Fig. 25.

Fig. 26 P. reichenbachiana

 Figure 26 shows other samples of the callus structure of P. fasciata. The callus (left) is for P. fasciata 'Rechela' which has a typical feature of P. fasciata. On the other hand, the callus (right) is for P. fasciata imported from Luzon. This callus structure has a unique feature in which there is no boss of tubercles in the posterior side but instead, a small bumpy glandulae. At the midport, small bifid denticulation is seen. A similar structure is observed in P. lueddemanniana v. delicata, P. pallens, P. corningiana, and P. zebrina. It is not clear whether it is one of the varieties or not.

 Based on such situations, it has been speculated that P. reichenbachiana might be a natural hybrid between P. fasciata and P. hieroglyphica. This has not been confirmed, though, because P. hieroglyphica is widely distributed in both Luzon and Mindanao islands; thus it may be possible for hybrids to develop; however, P. reichenbachiana is not found in Luzon, but only in Mindanao. P. fasciata and P. reichenbachiana are problematic species for identification, and there is room for further research; that is also the case for P. cornu-cervi.

Fig. 27 Callus Structures of P. fasciata (left) and P. reichenbachiana-like Species (right)

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P. Lueddemanniana and P. lueddemanniana v. Delicata

 The variant species have originated from a regional population which has a different morphology from the standard one in, e.g. shape, color and pattern. In recent years, the variations in color pattern on the perianth are not regarded as the determinant for the variety, but for the form. To categorize the species as a variety, it may need to have a certain genetic distance from the standard plant. P. delicata is now categorized as a variety of P. lueddemanniana. There are many differences in perianth, midlobe and callus structures between P. lueddemanniana and P. delicata, as shown in Figure 28. The sepals and petals of P. delicata are flatter and smaller than those of P. lueddemanniana, while the sepal and petal of P. lueddemanniana have a dissimilar form, and the lateral sepals have revolute margins. The midlobe is trowel-shaped in P. lueddemanniana, while oblong in P. delicata. The callus of P. lueddemanniana has a boss of dense tubercles in the posterior part (number 3 in the far right picture in Fig. 28), while P. delicata has no such posterior callus but a slight bumpy gland form. Considering such divergence, P. lueddemanniana and P. delicata would appear to be genetically distant, but this was not investigated in the past.

 Meanwhile, the callus of P. lueddemanniana is said to be biseriate. In the far right picture of the upper row in Fig. 28, it looks triseriate with three protrusions, marked as 1, 2 and 3. The ones labeled 1 and 2 are bifid denticulation, and 3 is a boss of dense tubercles. The denticulation of 2 might be grouped in with the posterior callus (3) if it was defined as biseriate.

 
Perianth 1
Perianth 2
Midlobe
Callus
Lueddemanniana
Delicata
Fig. 28 P. lueddemanniana and P. delicata 1
P. lueddemanniana
P. delicata
Fig. 29 P. lueddemanniana and P. delicata 2

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P. Violacea and Mentawai

 P. violacea and P. bellina are the most popular species in the genus Phalaenopsis. P. bellina is considered to be a sister species of P. violacea and inhabits Borneo, while P. violacea is distributed through the Malay Peninsula, Sumatra and Mentawai Island. Previously, P. violacea and P. bellina were regarded as the same species and were classified into three types (forms) - Borneo, Malaya and Sumatra. The Borneo form is now regareded as an independent species from P. violacea. A common assumption was that the Sumatra form came from a natural hybrid of the Malaya and Borneo forms. However, DNA analysis has verified that there is no difference between these forms. There are two reports on the genetic distance between P. violacea and P. bellina. One report states that P. bellina is genetically very distant far from P. violacea but is near to P. floresensis. The other one states that P. bellina is very near to P. violacea as a sister species. Such a difference in research results may depend on the reliability of the DNA samples.

 The problem with recent classifications of P. violacea is the morphological difference between P. violacea sumatra and P. violacea mentawai. According to DNA analysis, P. violacea mentawai has a certain genetic distance from P. violacea sumatra , and is assumed to be a new species developed from P. violacea in the Sumatra region. Table 5 lists the morphologies of P. violacea sumatra and mentawai.

Table 5 Morphology of P. violacea Sumatra and Mentawai
violacea sumatra
violacea mentawai
Perianth
Purple, red, coerulea
Pale purple
Flower Stalk
Flat, zigzag, shorter or equal to leaf
cylindrical, 50cm or more
Leaf
oblong-ovate
lanceolate, undulating
Root
thick
relatively thin

 
Flower
Inflorescences
leaf
Violacea
Mentawai
Fig. 30 P. violacea Sumatra and P. violacea Mentawai

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Sumatrana Complex

 P. sumatrana complex comprises P. sumatrana, P. zebrina and P. corningiana. As shown in Figure 31, P. sumatra is distributed in Borneo, Sumatra and Indochina, while P. zebrina is in Borneo, Sumatra and Palawan. P. corningiana is restricted to Borneo. The perianth of P. sumatrana is very similar to P. zebrina. P. sumatrana has many transverse brown bars and blotches on a pale yellow base. In contrast, P. zebrina has sparsely distributed transverse brown bars and blotches on a white base. The base color differs according to regional and individual differences. Thus, it may not be a definite determinant for identification. These species are said to have different fragrances.

 P. sumatrana and P. corningiana have the same pale yellow base color, but P. corningiana has dense reddish bars longitudinally arranged toward the apex, and the lateral sepal is nearly solid red in many species. P. corningiana is also found with sparsely distributed bars as shown on the "58 species" pages. Some people are of the opinion that P. sumatrana does not have longitudinal bars but always bars with a transverse alignment. However, there is a kind of P. sumatrana with longitudinal bars on the flower as shown in Fig. 31 (on the petals in far left picture). Therefore, the alignment of bars may not be a determinant of identification. Just as with P. fasciata and P. reichenbachiana, the identification of these species needs to be based on multiple features.

 A problematic issue arises on the callus structures of P. sumatrana and P. corningiana, which are defined as biseriate and uniseriate, respectively. Looking at the illustration by H. R. Sweet, the callus of P. corningiana is uniseriate. However, as shown in Fig. 31 (center of middle row), the protruding bifid denticulation at the posterior side on the midlobe is apparent, and furthermore, a small bump is observed behind it. It is hard to understand why this was defined as uniseriate. The callus structure of P. corningiana in the picture differs from the illustration by H. R. Sweet.

 It is interesting that the callus of P. corningiana is similar to P. zebrina. In contrast, the calluses of P. sumatrana and P. zebrina are quite different. The dense tubercles at the posterior callus in P. sumatrana are not found in P. zebrina. By all appearances, although the callus structures of P. sumatrana and P. zebrina, undoubtedly closely related as sister species, they must be classified as different species. Further investigation using considerable number of samples may be necessary to define the callus structures in these species.


P. sumatrana

P. corningiana

P. zebrina
 
Fig. 31 P. Sumatrana Complex

 Figure 31 also shows the anther caps of P. sumatrana and P. zebrina. It is not clear whether these are the definitive and common shapes of each species. Within a year or so (2010), this will be investigated in detail. It is said that the color, pattern, lip shape and scent of the flower have evoloved for the sole purpose of attracting the pollinator. If this assumption is correct, the structure of the anther cap might have evolved to adapt to the pollinator's body so it could be easily uncoupled from the column. In this sense, the structure of the anther cap may be important to understand the morphology of genus Phalaenopsis. It seems strange that such research hasn't been reported yet.

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Gigantea, doweryensis, maculataとCochlearis fuscata kunstleri

under construction

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