Available online at www.notulaebotanicae.ro
Print ISSN 0255-965X; Electronic ISSN 1842-4309
Not. Bot. Hort. Agrobot. Cluj 37 (1) 2009, 45-50
Notulae Botanicae Horti Agrobotanici
Cluj-Napoca
Application of Numerical Taxonomy to Lip Morphology in
the Genus Polystachya Hook (Orchidaceae) in Nigeria
Abayomi Ezekiel FALORUNSO, Adeniyi Akanni JAYEOLA
1)
Department of Botany, Obafemi Awolowo University, Ile-Ife, Nigeria: E-mail: afolorun@oauife.edu.ng
Abstract
SCLA, a numerical taxonomic method was applied to lip morphology in the genus Polystachya Hook. (Orchidaceae) in Nigeria. he
basic data matrix was prepared by coding for the presence or absence of the attributes of characters involved. he data were standardized
so that the values of a particular character were transformed into values ranging from zero to one. he SCLA showed that reproductive
characters are much better than the vegetative characters earlier used in the distribution of Polystachya species into their sections. In the
case of reproductive characters, more clusters were reported; this may be an indication of more sections in Polystachya than those earlier
reported.
Keywords: Polystachya species, cluster analysis, reproductive character, vegetative character, sections
Introduction
Polystachya is a genus of about 200 species, mostly
African but also occurring in Madagascar, Central and
South America and Asia (Cribb and Rasmussen, 2004).
Epiphytic, occasionally litophytic or terrestrial perennial
herbs (Croix, 1997). Hutchinson and Dalziel (1968) listed
ity seven species for west Tropical Africa, with Nigeria
having 24 species. New species from Tropical west and
South-Central Africa were described by Cribb and Croix
(1996) among which is P. anthoceros from Nigeria. Segerback (1983) described 104 species of orchids for Nigeria,
but Jayeola (1991) otherwise encountered over 400 species
for Nigeria.
he greatest challenge for the utilization of orchids in
Nigeria for both scientiic and aesthetic purposes is the
absence of simple identiication keys devoid of the usual
highly technical details of taxonomy ( Jayeola and folorunso, 2002). he existing keys are too technical to be
used by non-taxonomists such as foresters and horticulturists. Orchids are good lagship species and possible good
environmental indicators (Cribb and Croix, 1996). hey
contribute immensely to the national economy of Hawaii,
Malaysia, Singapore and hailand, whereas in Nigeria there
is little information about orchids. here is a great deal of
variation in the loral structure of Polystachya to the extent
that most species can be recognized on its basis ( Jayeola
and folorunso, 2002). he lowers are known to vary with
respect to colour, size and shape. Previous literature on the
use of lip show that they are of diferent sizes and shape
(Victoria, 1994; Luer, 1995; Catling, 1997). he lip otherwise known as the labellum is the relatively morphologically complex median inner perianth segment (petal)
that, in most orchids, is the primary attractant and landing
stage for pollinators, most commonly insects (Dafni, 1992;
Nilsson, 1992; van der Cingel, 1995; Rudall and Bateman,
2002).
Numerical taxonomy makes use of quantitative methods for the classiication of organisms into groups (Sokal
and Sneath, 1963). Most of the techniques used in numerical taxonomy to measure overall similarity between taxa,
as well as classifying them into groups are described by
Sokal and Sneath (1963). Morphometric techniques have
long been established as valuable tools for exploring the development, population diferentiation and systematics of
plants (Bookstein et al., 1985; wiens, 2000; forey and MacLeod, 2002; Jensen, 2003; Bateman and Rudall, 2006).
hese morphometric studies have typically employed between 20 and 50 quantiied characters, generally consisting
of a heterogenous mixture of metric (continuous), meristic,
scalar and presence / absence characters (Bateman and Rudall, 2006). Earnest study of the species of Polystachya has
become imperative, as some species are being threatened
by habitat loss, as for example Polystachya cooperi (Pollard
and Darbyshire, 2004). he purpose of the present study
is to describe the results of the use of single linkage cluster
analysis on the lip of the genus Polystachya in Nigeria and
to delimit the species into their traditional sections.
Materials and methods
Orchid lowers were obtained from two sources: fresh
lowers were obtained from the orchidarium, in situ, while
dried lowers were collected, ex situ, from the Botany Herbarium (IfE), Obafemi Awolowo University, Ile-Ife (Tab.
1). fresh lowers were dissected to expose the loral segments and dorsal petals or lips were excised. Preparations
of dried herbarium specimens for study required boiling
Folorunso, A. E. et al./ Not. Bot. Hort. Agrobot. Cluj 37 (1) 2009, 45-50
46
P_paniculata
P_paniculata_var_n
P_o_var_odorata
P_o_var_trilepidis
P_sp_novum
P_supfiana
P_albescens
P_subulata
P_ramulosa
P_laxiflora
P_stricta
P_calluniflora
P_golugensis
P_saccata
P_bifida
P_tesselata
P_modesta
P_fusiformis
P_orosun_var_novum
P_polychaete
P_o_var_orosun
P_adansoniae
P_cooperi
P_galeata
P_elegans
P_mukandaensis
P_dolichophylla
P_affinis
he phylogenetic relationships of the species of Polystachya is as shown in fig. 1. Six main clusters are observable at 0.5 similarity level. Cluster A, which is the largest
P_rhodoptera
P_alpina
Results and discussion
cluster encompasses many species of Polystachya covering
sections like Cultriformes, Aines, Grandilorae, Callunilorae, Isochiloidae and Dendrobiantae. he species are
P. dolichophylla, P. ainis, P. elegans, P. mukandaensis, P.
cooperi, P. galeata, P. odorata var. orosun, P. adansoniae, P.
orosun var. novum, P. polychaete, P. tessellata, P. modesta, P.
fusiformis, P. saccata, P. biida, P. callunilora, P. golugensis,
P. coriscensis P. camaridiodes, P. rhodoptera, P. alpina, P. ramulosa, P. laxilora, and P. stricta. Cluster B, comprised of
P. albescens and P. subulata, included in the section Caulescent. Clusters C and D belong to Polystachya section and
thus include two varieties of P. paniculata in addition to P.
odorata var. odorata and P. odorata var. trilepidis.
he characters used for the cluster analysis are representatives of both vegetative and reproductive characters
of the species Polystachya. he species of Polystachya here
listed are those reported for Nigeria (Hutchinson and
Dalziel, 1968) and the 8 species representing entirely new
records for the orchid lora of Nigeria ( Jayeola and folorunso, 2002).
he result of the cluster analysis revealed a lot of intraspeciic relationships among the species of Polystachya
P_coriscensis
P_camaridiodes
the lowers in ammonium chloride (NH4Cl) solution for
thirty minutes to revive the tissues of the lips for detailed
study under the microscope. he lips were then let to cool
for two to three hours ater which they were dissected in
the laboratory for examination. All data were scored from
a minimum of ten accessions for each species. he mean
values of these data were then coded (Tab. 2). A total of
twenty six characters were recorded for each OTU. he
basic data matrix of 26 x 32 was prepared by coding for
the presence or absence of the attributes of characters
involved. Data were standardized, so that the values of a
particular character were transformed into values ranging
from zero to one, using the method of Sokal and Sneath
(1963). Single linkage cluster analysis (SCLA) was performed to study the phylogenetic relationships of members of the genus.
0.96
0.84
0.72
Similarity
0.6
0.48
0.36
0.24
0.12
0
0
4
8
12
16
OTU
fig. 1. Phylogenetic relationships of the species of Polystachya studied
20
24
28
32
Folorunso, A. E. et al./ Not. Bot. Hort. Agrobot. Cluj 37 (1) 2009, 45-50
47
studied. Cluster A, which is the biggest of the clusters
is a conglomerate of ive sub-clusters. for instance P.
dolichophylla and P. ainis which have yellow lowers and
whose mid lobes are short, form the irst sub – cluster
comprised of P. elegans, P. mukandaensis, P. cooperi and P.
galeata. All these species have green lowers with the apex
of their lateral lobes acute. he third sub-cluster includes
P. odorata var. orosun, P. andansoniae, P. orosun var. orosun,
P. polychaete, P. tesselata, P. modesta, P. fusiformis, P. saccata and P. biida. hese are the species with white lowers
together with purple markings. he fourth sub-cluster includes species like P. callunilora, P. golugensis, P. coriscensis,
P. camaridiodes and P. rhodoptera with pale green lowers.
he ith sub-cluster comprises of P. alpina, P. ramulosa, P.
laxilora, P. stricta and they are largely white with yellow
colour that is tinged with pink. Cluster B comprises P. albescens and P. subulata.
hey have greenish white lowers and belong to the
Caulescent section. he species in cluster C are with whitish yellow lowers tinged with rose. hose in cluster D have
orange lowers with red markings.
Conclusions
his work has shown that reproductive characters are
much better than the vegetative characters earlier used by
Kraenzlin (1926) in the distribution of Polystachya species
into their sections. with the use of reproductive characters more clusters have been reported which also indicate
the occurrence of many more sections in Polystachya than
those earlier reported.
References
Bateman, R. M. and P. J. Rudal (2006). Evolutionary and
Morphometric Implications of Morphological Variation
Among Flowers Within an Inlorescence: A case-study Using
European Orchids. Annals of Botany. 98:975-993.
Bookstein, F. L., B. Chernof, R. Elder, J. Humphries, G. Smith and
R. Strauss (1985). Morphometrics in evolutionary biology.
Philadelphia, PA: Philadelphia Academy of Natural Sciences.
Catling, P. M. (1997). he taxonomic status of Fernald’s ragged
Orchid of Newfoundland, Plantanthera lacera var. terraenovae. Lindleyana. 12(2):79-88.
Van, der Cingel (1995). An atlas of orchid pollination: European
orchids. Rotterdam: Balkema.
Cribb, P. J. and I. Croix (1996). New Polystachya species
(Orchidaceae) from tropical west and south-central Africa.
Kew Bulletin. 51(3):571-577.
Cribb, P. and J. Rasmussen (2004). Field guide to the Ethiopian
Orchids.
Croix, I. (1997). African Orchids in the wild and cultivation.W3
Tropicos, Kew Monocot list, IPNI.
Dafni, A. (1992). Pollination ecology. Oxford: Oxford University
Press.
Forey, P. and N. MacLeod (2002). Morphology, shape and
phylogenetics. In: Taylor and Francis (Eds.). Systematics
Association Special Vol. 64. London.
Hutchinson, J., J. M. Dalziel (1968). Flora of West Tropical Africa.
3(1):215-225.
Jayeola, A. A. (1991). Computer-aided taxonomic study of the
angraecoid orchids of Nigeria and Cameroon. Obafemi
Awolowo University. Ph.D. hesis. 16-91.
Jayeola, A. A. and A. E. Folorunso (2002). Floral morphology of
the Polystachya Hook.
(Orchidaceae) in Nigeria. Nigerian Journal of Horticultural
Science. 6:70-75.
Jensen, R. J. (2003). he conundrum of morphometrics. Taxon.
52:663-671.
Kraenzlin, F. (1926). A monograph of the genus Polystachya.
Feddes Repertorium; Beih. Vol.39.
Luer, C. A. (1995). A new species of Pleurothallis (Orchidaceae)
from French Guyana Bulletin du Museum. National d’ Histoire
Naturelle section B. Adansonia Botanique. Phytochimie.16(24):231-233.
Nilsson, A. (1992). Orchid pollination biology. Trends in ecology
and Evolution. 7:255-259.
Pollard, B. J. and I. Darbyshire (2004). Polystachya cooperi. 2006
IUCN Red List of hreatened species.
Rudall, P. J. and R. M. Bateman (2002). Roles of synorganisation,
zygomorphy and heterotopy in loral evolution: the
gynostemium and labellum of orchids and other lilioid
monocots. Biological Reviews. 77:403-441.
Segerback, L. B. (1983). Orchids of Nigeria. A. A. Balkema
Publishers. A member of Swets and Zeitlinger Publishers.
Netherlands.
Sokal, R. R. and P. H. A. Sneath (1963). Principles of Numerical
Taxonomy. W.H. Freeman and Co. San Francisco, Publishers.
Victoria, S. (1994). Bletia greenwoodiana (Orchidaceae), a new
species from Durango, Mexico. Brittonia, 46(3):208-210.
Wiens, J. J. (2000). Phylogenetic analysis of morphological data.
Washington, DC: Smithsonian Institution.
.OTU coded values of the
species of Polystachya
Lipcelled
P_galeata
P_stricta
P_elegans
P_golugensis
P_o_var_odorata
P_o_var_orosun
P_o_var_trilepidis
P_mukandaensis
P_adansoniae
P_orosun_var_novum
P_alpina
P_paniculata
P_callunilora
P_dolichophylla
P_polychaete
P_coriscensis
P_camaridiodes
P_albescens
P_paniculata_var_novum
P_cooperi
P_supiana
P_sp_novum
P_saccata
P_biida
P_rhodoptera
P_afinis
P_tesselata
P_laxilora
P_subulata
P_fusiformis
P_modesta
P_ramulosa
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
1
0
0
0
48
Lip-pubes- Lip-apex- Lip-apexcent
acute
depre-ssed
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
1
0
1
0
1
0
0
0
0
1
1
1
0
0
0
0
0
0
0
1
1
1
0
0
1
1
1
1
1
0
0
0
0
0
0
0
0
1
0
0
1
Lipmarginentire
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
Apexmidlobeacute
1
1
1
1
0
1
0
1
1
1
1
1
1
0
1
1
1
1
1
1
1
0
0
1
1
1
1
1
1
1
0
1
Marginincurvedbelow
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
Venationnetwork
Liplarge
Lipsmall
Lipmoderate
Lipleshy
Lipwithoutcallus
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
1
0
1
1
0
0
1
1
0
1
0
0
0
0
0
1
0
0
0
0
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
1
1
0
1
0
0
0
0
1
0
0
1
.OTU coded values of the
species of Polystachya
Lipclawed
P-galeata
P-stricta
P-elegans
P-golugensis
P-o-var-odorata
P_o_var_orosun
P_o_var_trilepidis
P_mukandaensis
P_adansoniae
P_orosun_var_novum
P_alpina
P_paniculata
P_callunilora
P_dolichophylla
P_polychaete
P_coriscensis
P_camaridiodes
P_albescens
P_paniculata_var_novum
P_cooperi
P_supiana
P_sp_novum
P_saccata
P_biida
P_rhodoptera
P_ainis
P_tesselata
P_laxilora
P_subulata
P_fusiformis
P_modesta
P_ramulosa
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
1
0
1
1
0
0
0
0
0
0
0
0
0
0
Lip-notLip-transwell deveparent
loped
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
Lippunctate
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
1
0
0
0
1
0
0
1
0
0
1
1
0
0
0
LipLip-lobed- lobed-at
atmiddle
lower
part
0
0
0
0
1
0
0
0
0
0
0
1
0
1
0
1
1
1
1
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
Lipaslongaswide
Flowerswhite
Flowers-red
Flowersgreen
Flowersyellow
Flowerstinged-pink
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
1
0
0
0
1
0
1
0
1
0
0
1
0
0
1
0
1
0
1
0
0
1
0
1
0
0
1
1
0
0
0
1
1
0
1
1
1
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
1
1
0
0
0
1
0
0
0
0
0
0
1
1
1
1
0
1
0
0
0
0
0
0
1
0
1
1
1
0
0
1
1
1
1
0
1
1
0
0
0
0
0
1
1
1
1
0
0
0
1
0
0
0
1
1
1
1
0
0
1
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
Flowers-49
greenishyellow
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
Tab. 1. Representative Specimens of the Polystachya species in Nigeria.
1.
50
2.
3.
P. dolichophylla Schltr. Keay 39, Nindam F.R. Zaria.
P. ainis Lindl. Burtt B27, Abeokuta to Ijebu Ode
FHI 24666 Keay Usenigbe F.R. Sapoba
FHE 22714 Keay, Owo, Ondo Prov.
Talbot 3776, Ohbe, Degema, Randole Barter 1863, Onitsha
P. elegans FHI 25376 Keay Sankwala, Ogoja prov.
P. mukandaensis De Willd. FHI 22410 Keay, Olokemeji F.R. Abeokuta
Tab. 1
P. golungensis Rchb.f. King 22, Kogin Delli, Zaria Prov.
17.
Olokemeji F.R. Keay. FHI 22712 Abeokuta, Akure F.R. Keay 28294 Ondo
18.
P. coriscensis Rchb.f. Sanford WS/185/66, Benin Prov. Sapoba
P. camaridiodes Summerh. Wright 101, Calabar
4.
a. homas 1902 Onika Ohene FHI 6128 Janes, Onitsha Talbot 861, Oban
19.
5.
Cooper 80, Obudu Scarp, Ogoja
P. cooperi Summerh. FHI 15190; WS 5458 IFE
20.
P. rhodoptera Rchb.f Talbot 903 Oban, Sapoba Kennedy 1917
21.
P. alpina Lindl. Talbot 835, Oban Niagi
22.
P. ramulosa Lindl. Sanford WS/184/66 Benin Prov. Sapoba Talbot 3647, Oban Degema
P. laxilora Summerh. FHI 1366, WS/1521/65 Idanre Hills
P. galeata Brenan 9116, Okomu F.R. Benin
6.
7.
8.
9.
10.
11.
12.
13.
a. FHI 227738a Keay Wright 128, Ajagbodudu, Sapele
b. Talbot 3725 Degema Cooper 82, Ogoja Plain FHI 25304 Keay Coombe 151
FHI 22429 Keay, Eleyele, Ibadan
P. odorata var. orosun Sanford 4135, Orosun peak Idanre
P. adansoniae Rchb. F. FHI 40169 Onochie, Niger Prov. Gwari, Bornu
a. Sanford WS 216a, 221/66 Boyogbe, 5 Benin Province; FHI 25317 Ejiofor
b. Ogoja Province FHI 22438 Keay Okomu F.R., Benin
Prov. Talbot 867,927 Kundene Obari
P orosun var. novum. Sanford 4123, Orosun Peak, Idanre
P. polychaete Kraenzl. Talbot 927, 928 Oban
P. tesselata Lindl. King 54, Kushanka, zaria Prov.
Elliot 80, FHI 33190 Onochie, Stubbs Creek F.R. Eket FHI 13348 Keay, Calabar
P. modesta Rchb. F. Cole 13, Zaria king 31, Mongu F.R. Plateau Province
FHI 22443 Keay, Sanga River F.R., Zaria, WS/1848765 Agodi Gate, Ibadan.
P. fusiformis ( hou.) Lindl. IFE WS/5668
23.
24.
25.
26.
Daramola FHI 29809, Talbot 918 Oban
P. stricta Rolfe Winbush 611; King 350 Manbia Plateau
P. albescens Ridl. WS 423 IFE
P. subulata Finet FHI 22898 Keay Ribaka F.R. Zaria Prov.
FHI 37667 Keay, Mangu F.R. Plateau Prov.
Summerhayes 91, Kaliy Zankwa Zaria, Rchb.f.
27.
P. sp. novum WS/443/66 sapoba F.R.
28.
P. supiana Schltr. Talbot 440, Kwa F. Oban, FHI 3306, Eket; IFE WS/491/66. Sapoba
P. saccata V.S.Summerh. 90 Kagarko, Zari FHI 37620
Keay and Janes Gimi River F.R. Zaria
29.
P. odorata var. odorata Barter 1483, Onitsha 0.5cm
Kennedy 1686 Sapoba Richards 3409, Shasha F.R. Ijebu FHI 33176 Onochie
Orukini Unyene, Eket FHI 33209 Onochie Lower Enyong F.R. Itu
FHI 37639 Keay and Janes Sanga River F.R. Zaria
30.
14.
P. odorata var. trilepidis Summerh. FHI 20729bJanes, Mt. Orosun,
Idanre, Ondo.
FHI 21559 Keay and Onochie FHI 3374 Symington.
15.
P. biida Summerh. IFE WS/5990, WS/199/66 sapoba F.R. Obudu Plateau Ogoja
31.
16.
P. callunilora Kraenzl. WS/3541 IFE
32.
P. paniculata (SW.) Rolfe; Keay, FHI 25371, Akure F.R.; Ejiofor FHI 24653
homson FHI 1509, Amedzofe; Ngogi, FHI 15341
P. paniculata (SW.) Rolfe subsp. novum