preliminary chloroplast dna studies in the flowering plant family martyniaceae (order lamiales)

Arizona-Nevada Academy of Science

Preliminary Chloroplast DNA Studies in the Flowering Plant Family Martyniaceae (OrderLamiales)Author(s): Raul GutierrezSource: Journal of the Arizona-Nevada Academy of Science, Vol. 40, No. 1 (2008), pp. 105-110Published by: Arizona-Nevada Academy of ScienceStable URL: http://www.jstor.org/stable/27641776 .

Accessed: 15/06/2014 21:55

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp

.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact [email protected].

.

Arizona-Nevada Academy of Science is collaborating with JSTOR to digitize, preserve and extend access toJournal of the Arizona-Nevada Academy of Science.

http://www.jstor.org

This content downloaded from 185.44.77.28 on Sun, 15 Jun 2014 21:55:11 PMAll use subject to JSTOR Terms and Conditions

http://www.jstor.org/action/showPublisher?publisherCode=anas

http://www.jstor.org/stable/27641776?origin=JSTOR-pdf

http://www.jstor.org/page/info/about/policies/terms.jsp


Preliminary Chloroplast DNA Studies in the Flowering Plant Family Martyniaceae (Order Lam?ales)

Raul Gutierrez, School of Life Sciences, Arizona State University, PO Box 874601, Tempe, AZ

85287-4601

Abstract

Acceptance of the plant group Martyniaceae, which contains the genus Proboscidea (commonly known as devil's

claws or unicorn plants), as a distinct family has long been questioned. Previously placed in the family Pedaliaceae,

commonly known as the Sesame family, the Martyniaceae have been allied to numerous other families within the order

Lam?ales. For this study, sequences of chloroplast DNA from the ndhF and rpsl6 regions were used (1) to demonstrate

the monophyly of the Martyniaceae and (2) to identify the closest living relative to the group. Preliminary data suggest that the Martyniaceae are distinct from the Pedaliaceae, and that the Verbenaceae, commonly known as the Verbena or

Vervain family, may be the closest living relative to the family Martyniaceae. If the Martyniaceae and Verbenaceae are

sister groups and most closely related to each other, then there are interesting biogeographic implications that can be

suggested from the results. The Verbenaceae are primarily from arid areas of South America, which happens to be one

of the centers of diversity for the family Martyniaceae. If the Verbenaceae are the most closely related group of plants to the family Martyniaceae, then it can be suggested that the Martyniaceae may have had an origin in South America, then dispersed into and diversified in North America.

Introduction The family Martyniaceae consists of 13-16

species in the New World in five genera that are

closely allied to, or confamilial with, Pedaliaceae

(commonly known as the sesame family). It is

indigenous to the New World and found primarily in subtropical and warm temperate areas from the

United States (mostly from Louisiana to California) southward through Central America to Uruguay and

Argentina (Thieret 1977). A few species have been

introduced to other parts of the world as ornamentals, often escaping cultivation and quickly becoming

weeds (Thieret 1977). The family Martyniaceae is characterized by

having a unilocular and bicarpellate ovary with par ietal placentation (Thieret 1977). Glandular hairs,

composed of one- to several-celled uniseriate stalks

and a several-celled head, cover the surface of the

plant giving the plants a sticky texture (Thieret

1977). The most outstanding characteristic of the

family is the presence of woody, beaked fruits

(Thieret 1977). As these fruits mature, the fleshy

exocarp senesces and sloughs off, revealing the

woody endocarp underneath (Thieret 1977). These

fruits dehisce longitudinally along abiaxal and

adiaxal sutures and produce two sharp-pointed, hooked horns when the beak splits (Thieret 1977). The largest genus in the family, Proboscidea, is

common in arid and semiarid areas of the United

States and Mexico. They are commonly known as

devil's claws or unicorn plants, and in addition to

the generic name Proboscidea (the name of the

mammalian order containing elephants and mam

moths), refer to the elongated rostrum on the fruits.

De Jussieu (1789) and Kunth ( 1822-1825) have included the Martyniaceae within the Bignoniaceae, a family that includes desert willows (Chilopsis

linearis) and yellow bells (Tecoma stans). Martyni aceae differs from the family Bignoniaceae by the

presence of parietal placentation and wingless seeds.

Pollen morphology (Martin and Drew 1970, Bret

ting and Nilsson 1988) suggests that Martyniaceae

may have a close affinity to Bignoniaceae.

Cronquist (1981) has rejected the Martyniaceae as

a distinct family, considering it conceptually more

useful to include the Martyniaceae within the Peda

liaceae. Plants in both families have herbaceous

habits, specialized trichomes on their surfaces, and

pod-shaped fruits armed with hooks, horns, or

prickles (Cronquist 1981). Thus, Cronquist recog nized the two groups as the subfamilies Pedalioi

deae and Martynioideae within the family Pedalia

ceae (along with a third subfamily, Trapelloideae).

Stapf ( 1895) was the first to combine the genera

Martynia, Craniolaria, and Proboscidea into anew

family, the Martyniaceae (Hevly 1969, Bretting

1981). Most systematic treatments since then,

including those of Van Eseltine (1929) and Bretting

(1981), have followed Stapf s treatment and retained

Martyniaceae as a distinct family. Most recently, the

Angiosperm Phylogeny Group (APG) has endorsed

retention of the Martyniaceae as a distinct family within the order Lamiales (APG II 2003). In addi

tion to the families mentioned above (Bignoniaceae and Pedaliaceae), the order as circumscribed by

APG II includes the families Acanthaceae (acanthus

family), Lamiaceae (mint family), and Verbenaceae

(verbena family). All of these families have pentam

Gutierrez, R. 2008. Preliminary chloroplast DNA studies in the flowering plant family Martyniaceae (Order

Lam?ales). Journal of the Arizona-Nevada Academy of Science40(1 ): 105-110.



106 Preliminary Chloroplast DNA Studies in Martyniaceae + Gutierrez

erous fused corollas, bicarpellate ovaries, and

epipetalous stamens (APG II 2003). A recent study of the family Martyniaceae,

utilizing the Internal Transcribed Spacer region of nuclear ribosomal DNA (nrDNA ITS), included four of the five genera and 12 taxa

(Gutierrez 2002). The closest relative to the

Martyniaceae was a clade consisting of mem

bers of the Lamiaceae, though bootstrap values failed to support this relationship (Gutierrez 2002). Within the family Martyniaceae, two

distinct clades are formed roughly correspond ing to the distribution of the genera in North and South America (Fig. 1 ). The predominantly

North American genera of Martynia and Proboscidea are placed together, as are the pre

dominantly South American genera of Craniolaria and Ibicella (Fig. 1). The place ment of the taxa within Proboscidea into three

infrageneric groups by Bretting (1981) is supported. These three groups correspond to (1)

Proboscidea sabulosa, (2) P. altheaefolia, and (3) the "weedy" annual species (P. triloba, P. pasi

flora, P. louisianica, and P. spicatd) and their corre

sponding subspecies (Fig. 1). Of these groups, P. sabulosa is either shown as sister to the other two

groups of Proboscidea, or it is shown to form a

polytomy with Martynia annua and the rest of the taxa in Proboscidea (as shown in Fig. 1). To com

plete the sampling of the taxa in the Martyniaceae, eight additional taxa need to be sequenced, inclu

ding three in the genus Proboscidea. Prior to the ITS study, taxonomic works utilized

only morphological data and focused on the genus Proboscidea (Bretting 1981). Additional chloroplast DNA genes employed here address the following questions: (1) Do the Martyniaceae belong in the

Pedaliaceae or are the Martyniaceae monophyletic and distinct from the Pedaliaceae and thus warrant

separate family status? (2) If the Martyniaceae are

segregated from the Pedaliaceae, are they sister to

the Pedaliaceae, or is another group within the

Lamiales, such as the family Verbenaceae or Lamia

ceae, more closely related?

Materials and Methods For the purpose of studying the placement of

Martyniaceae within the Lamiales, the chloroplast DNA regions ndhF and rpsl6 were used. These

regions have previously been used to study the order Lamiales (Olmstead et al. 2000, Bremer et al. 2002) and groups within the order (Acanthaceae, Mank telow et al. 2001; Buddlejaceae, Oxelman et al.

1999; Antirrhineae, Ghebrehiwet et al. 2000; Olea

ceae, Wallander and Albert 2000; Scrophulariaceae,

70

63

100:

100

69

68 56

Proboscidea parviflora hohokamiana

Proboscsdea parviflora parviflora

Proboscidea parviflora sircaloensis

Proboscsdea Iouisianica fragrans

Proboscidea spicata

Probosadea inloba

Proboscidea Iouisianica Iouisianica

Probosadea aiihaeifoila1

Martyr? a annua North America

Proboscidea sabulosa'

Cram?tari-a annua

Ibicella lutea

Sotttii Amenai

.. Mentha

Figure 1. Phylogenetic tree constructed using nrDNA ITS data.

Adapted from Gutierrez (2002). Bootstrap values are placed above branches supported by an analysis using 1000 replicates.

Olmstead and Reeves 1995, Oxelman et al. 2005),

though the Martyniaceae and related families were

often under sampled. Taxa from the Martyniaceae and allied families were included in this analysis. A

query of GenBank (www.ncbi.nlm.nih.gov) pro duced several sequences of ndhF and rpsl6 that

were used as a starting point for this study (Appen dix I). The outgroup, Leucophyllum, has been

chosen from the Scrophulariaceae s.s. (Olmstead et

al. 2001, Oxelman et al. 2005), as this family forms a sister relationship with the rest of the core Lam

iales. A complete of list of taxa to be used for the

study and their GenBank Accession Numbers are

included in Appendix I.

Genomic DNA samples was extracted from fresh tissue, silica dried samples, or herbarium

specimens using a Qiagen DNeasy? Plant Mini Kit

(Qiagen Inc., Valencia, CA). Two PCR primer sequences for rpsl6 (rpsl6_F and rpsl6_2R) were

taken from Bremer et al (2002), while PCR of the entire ndhF gene was attempted using the primer pairs 1*/1350R and 1201/2112R (Oxelman and Reeves 1995, Olmstead and Sweere 1995, Oxelman et al 1999). The first set of primers were to be used for the first half of the ndhF gene, but were not very successful in amplification and no sequences were

generated from this portion of ndhF for this anal

ysis. The second set of primers (1201/2112R) was

successful in amplifying and sequencing the second half of the ndhF region. Thus, only the second half of the ndhF region was used in the analysis.

A total of 25 sequences of rpsl6 and 15

sequences of ndhF were generated using the same

primers employed in the PCR amplifications. Of

these, only 14 sequences of rpsl6 and 9 sequences of ndhF were used in this analysis. The rest of the



Preliminary Chloroplast DNA Studies in Martyniaceae + Gutierriez 107

sequences were not included because the taxa they

represented lacked sequences from the other region.

Additionally, due to the problems encountered try

ing to amplify the first half of the ndhF region, only the second half of this region was used in the

analysis. Sequences from the rps 16 region averaged about 950 bases, while sequences from the second

half of ndhF region averaged 1,100 bases. The total

length of the ndhF region averages 2,200 bases, so

the exclusion of half of this region represented a

significant loss of possible informative characters.

Sequences were aligned using CLUSTALW

(Thompson et al. 1994) and placed into a data

matrix that included 40 taxa and a total of 2,100 characters. Trees were constructed using parsimony

using PAUP* 4.0b 10 (Swofford 2002) using default settings, and a bootstrap analysis was run with 1,000

replicates (Fig. 2). The bootstrap is a type of statis

tical analysis of the data that is used to test con

fidence in the data.

]i)U

100

~~~ fclsr-ofra

? Coig?a

? Lafruri ? Lancea

~ """*""~ ?gala's

-Prr^s -rie<n cnaena

-D places ~?~ - Cs1s pa

_^-"%song -Tesonera

-PaUcwita ?? 5c*i eqeiia

; .- ? Lar tara -Vemei?

<-Yaryn a

t-Pro so se cea - Osno ana

.?~? itnc* ta -Jacaranda

i ,-Cnsssarcra *--acanthus i-A? cennia rra-ra u""~ f>* csni?a ge1^ ".

- RJg |,a -J^s? esa

" ?-? Arerag^pr-is -Bar er a

- ? -- Tnutoe'gia

-Ce'a'o'neca "-,-_ Sesa??UTi ?ndisu?T:

- - - SesaTuTi racS=a?ui?t ??~?- Ha'pago^ytyir;

-i tarop? un

i aii?iaccac

Bigiiotiiaeeae

Verben?ceas

Martyniaceae

?canthaceae

Pedal ?accac

Figure 2. Strict consensus of the two most parsimonious

trees constructed using ndhF and rps 16. Bootstrap values

are placed above branches supported by an analysis using

1,000 replicates.

Results and Discussion Parsimony analysis of the data matrix yielded

two equally most parsimonious trees. A strict consensus tree was constructed and is shown in Fig ure 2 with bootstrap values, if any, above the

branches. The bootstrap values represent the per

centage of resampled matrices in which that rela

tionship is retained. Higher bootstrap values (>70%)

represent well-supported lineages, whereas lower

bootstrap values (50-70%) fail to provide enough

support for that relationship, and that relationship may collapse into an unresolved polytomy (if

<50%). The bootstrapping analysis performed on

this data set yielded few supported relationships, with most supported relationships being within

well-defined families such as the Verbenaceae,

Pedaliaceae, Acanthaceae, and Lamiaceae. Deeper

relationships within the Lamiales are not supported, and this lack of support will cause the branches to

collapse into one large polytomy consisting of

several unresolved lineages. This is a problem encountered in other studies (Olmstead et al. 2001,

Bremer et al. 2002, Oxelman et al. 2005) in which a radiation of families occurs without any resolution

in relationships between these families.

The monophyly of many of the larger families

in the Lamiales are demonstrated. This includes the

monophyly of the Martyniaceae, though bootstrap

support is very low. Though its segregation from the

Pedaliaceae is not supported due to the lack of boot

strap support in the deeper branches of the phy

logeny that results in an unresolved polytomy, the

high bootstrap value of the Pedaliacean lineage

(100%) does provide some support for the separa tion of the two lineages into different families.

Martyniaceae is placed as sister to the Verbena ceae in both parsimonious trees (and thus the strict

consensus), though bootstrap values do not give any

support for this relationship. Still, placement of

these two families as sister taxa agrees with pre vious trees constructed by Olmstead et al. (2001) and Oxelman et al. (2005), though these trees also

lack significant support (both <50%). A recent anal

ysis of taxa of the family Verbenaceae by Olmstead

(pers. comm.) using nuclear and chloroplast DNA

suggests that the family Verbenaceae evolved in

South America and dispersed northward through Central America and the Caribbean into North

America. If the sister relationship between Marty niaceae and Verbenaceae holds up, then it may be

suggested that the Martyniaceae also had an origin in South America and may have dispersed and

diversified in arid and semiarid areas of North

America.



108 Preliminary Chloroplast DNA Studies in Martyniaceae + Gutierriez

Aside from the two works using molecular data

(Olmstead et al. 2001, Oxelman et al. 2005), the sister relationship between Verbenaceae and Marty niaceae has never been suggested. It may be useful to find morphological characters that will further

support this relationship, as there is no morpholog ical data at this time that can support the tenuous

nature of the hypothesized relationship. Additional support may also come from the

inclusion of more complete taxon sampling from the

groups involved. The Verbenacean taxa included in

this study (Fig. 2, Appendix I) are from genera that

span North and South America (Aloysia and

Verbena) or are primarily Central American and

Caribbean (Stachytarpheta). Future analyses should

include more taxa from the Verbenaceae, especially those genera that are entirely South American and

not included in this analysis. Additionally, more

taxa of South American genera of the family Lamiaceae should be included, as the boundary between the Verbenaceae and Lamiaceae is not well

understood. Future directions may also include the addition

of more characters in the data matrix. Difficulty in

the amplification and sequencing of the first half of

the ndhF region precluded the inclusion of these

data in the analysis. It is proposed that new primers be screened for their utility in the amplification and

sequencing of this region so that an additional 1100

characters can be included in the data matrix. These

additional characters may help to provide additional

support through bootstrapping analysis, giving

greater credence to the relationships that are hypoth esized in future analyses.

Acknowledgments The author thanks curators Les Landrum, Arizona

State University Herbarium and Richard Worthington, University of Texas-El Paso Herbarium for providing plant material used for the extraction of DNA, and M. F.

Wojciechowski for lab supplies, lab space, support, and

helpful comments on the manuscript. A Grant-in-Aid of Research from the Arizona-Nevada Academy of Science

provided funding for this work.

I ITFRATIIRFClTPn APG IL 2003. An update of the angiosperm phy

logeny group classification for the orders and

families of flowering plants: APG II. Botanical

Journal of the Linnaean Society 141:399-436.

BAILLON, H. 1888. Gesn?riac?es. Historum PI.

10:59-112.

Bremer, B., K. Bremer, N. Heidari, P, Erixon, R.

G. Olmstead, A. A. Anderberg, M. Kal

lersjo, and E. Barkhordarian. 2002. Phylo

genetics of Asterids based on 3 coding and 3

non-coding chloroplast DNA markers and the

utility of non-coding DNA at higher taxonomic

levels. Molecular Phvlogenetics and Evolution

24:274-301.

Bretting, P. K. 1981.,4 Systematic and Ethnobo

tanical Study o/Proboscidea and Allied Genera

of the Martyniaceae. Ph.D. thesis, Indiana Uni

versity, Bloomington. Bretting, P. K., and S. Nilsson. 1988. Pollen

morphology of the Martyniaceae and its sys tematic implications. Systematic Bot anv 13:51

59.

Cronquist, A. 1981. An Integrated System of

Classification of Flowering Plants. Columbia

University Press, New York. 1262 pp. DEJUSSIEU, A. L.. 1789. Generaplantarum secun

dum ordines naturales disposita. Paris. 498 pp.

Ghebrehiwet, M. B. Bremer, and M. Thulin.

2000. Phylogeny of the tribe Antirrhineae

(Scrophulariaceae) based on morphological and

ndhF sequence data. Plant Systematics and Evo

lution 220:223-239.

GUTIERREZ, Jr., R. 2002. A Molecular Phylogeny of the Family Martyniaceae (Order Lam?ales) based on nrDNA Internal Transcribed Spacer

Sequences. Master's Thesis, University of Texas

at El Paso, El Paso, Texas.

HEVLY, R. H. 1969. Nomenclatural history and

typification of Martynia and Proboscidea

(Martyniaceae). Taxon 18:527-534.

KUNTH, C. S. 1822-1825. Synopsis plantarum, quas in itinere adplagam aequinoctialem orbis novi,

collegerunt AI. De Humbolt et Am. Bonpland. 4 vols. Paris.

Manktelow, M., L. A. Mcdade, B. Oxelman, C.

A. Furness, and M. J. Balkwill. 2001. The

enigmatic tribe Whitfieldieae (Acanthaceae): Delimitation and phylogenetic relationships based on molecular and morphological data.

Systematic Botany 26:104-119.

Martin, P. S., and C. M. Drew. 1970. Additional

scanning electron photomicrographs of South

western pollen grains. Journal of the Arizona

Academy of Science 6:140-161.

McDade, L. A., and M. L. MOODY. 1999. Phylo

genetic relationships among Acanthaceae: evi

dence from noncoding trnL-trnF chloroplast DNA sequences. American Journal of Botany 86:70-80.

Olmstead, R. G, C. W. Depamphilis, A. D. Wolfe, N. D. Young, W. J. Elisens, and P. A. Reeves.

2001. Disintegration of the Scrophulariaceae. American Journal of Botany 86:70-80.

Olmstead, R. G, R. K. Jansen, R. J. Kim, and S. J.

Wagstaff. 2000. The phylogeny of Asteridae



Preliminary Chloroplast DNA Studies in Martyniaceae + Gutierriez 109

s.l. based on chloroplast ndhF sequences. Molec

ular Phylogenetics and Evolution 16:348-361.

Olmstead, R. G, and P. A. Reeves. 1995. Evi

dence for the polyphyly of the Scrophulariaceae based on chloroplast rbcL and ndhF sequences. Annals of the Missouri Botanical Garden 82:176-193.

OLMSTEAD, R. G, AND J. A. SWEERE. 1994.

Combining data inphylogenetic systematics: an

empirical approach using three molecular data sets in the Solanaceae. Systematic Biology 43:467-481.

Oxelman, B., M. Backlund, and B. Bremer. 1999. Relationships of the Buddlejaceae s.l.

investigated using parsimony jackknife and branch support analysis of chloroplast ndhF and

rbcL sequence data. Svstematic Botany 24:164 182.

Oxelman, B., P. Kornhall, R. G Olmstead, and B. Bremer. 2005. Further disintegration of the

Scrophulariaceae. Taxon 54: 411-425.

Shwarzbach, A. E., and L. A. McDade. 2002.

Phylogenetic relationships of the mangrove

family Avicenniaceae based on chloroplast and nuclear ribosomal DNA sequences. Svstematic

Botany 27r: 84-98.

STAPF, O. 1895. Pedaliaceae and Martyniaceae.

Pp. 264-269 in A. Engler and K. Prantl, Die Naturlichen Pflanzenfamilien. Teil IV, Abt. 3b.

SWOFFORD, D. L. 2002. PAUP*: Phylogenetic Analysis Using Parsimony (* and other meth

ods), version 4.0b 10. Sinauer, Sunderland, MA.

Thieret, J. 1977. The Martyniaceae in the south eastern United States. Journal of the Arnold

Arboretum 58: 25-39.

Thompson, J. D., D. G Higgins, and T. J. Gibson. 1994. Clustal W: Improving the sensitivity of

progressive multiple sequence alignment through sequencing weighting, position specific gap penalties and weight matrix choice.

Nucleic Acids Research 22:4676- 4680. Van Eseltine, G P. 1929. A preliminary study of

the unicorn plants. New York State Agricultural Experiment Station Technical Bulletin 149:1-41.

Wallander, E., and V. A. Albert. 2000. Phy logeny and classification of Oleaceae based on

rpsl6 and trnL-F sequence data. American Journal of Botany 87:1827-1841.

Appendix I. List of taxa used in the study. Data listed

includes family, taxon, ndhF Genbank Accession

number, and rpsl6 Genbank accession number. NEW designates sequences generated for this study and not yet accessioned in GenBank.

Acanthaceae: Acanthus montanus T. Anderson,

AJ429115, DQ059146; Andrographis paniculata Nees, NEW, NEW; Aphelandra squarrosa Nees,

AJ249405, DQ059200; Avicennia germinans (L.) Steams, NEW, NEW; Avicennia marina (Forssk.)

Vierh., AJ429116, AJ431038; Barleria micans

Nees, *, NEW; Barleria prionitis L., U12653, *; Crossandra nilotica Oliver, U12656, *; Crossandra

strobilifera (Lam.) Benoist, *, DQ059176; Elytraria crenata Vahl, U12657, *; Elytraria imbricata

(Vahl) Pers., *; NEW; Justicia adhotada L., *,

DQ059214; Justicia americana (L.) Vahl, U12663, *; Ruellia ciliosa Pursh, U12664, *; Ruellia humilis

Nutt., *, AF482538; Thunbergia alata Bojer ex

Sims, U12667, AJ609131.

Bignoniaceae: Catalpa sp., L36397, *; Catalpa speciosa Warder ex Engelm., *, AJ609197; Jacaranda mimosifolia D. Don, NEW, AJ431039;

Macfadyena unguis-cati (L.) A. H. Gentry, NEW,

NEW; Tecoma stans (L.) H.B.& K., AF130145, NEW; Tecomeria capensis (Thunb.) Spach, DQ222642, NEW.

Lamiaceae: Congea tomentosa Roxb., U78689,

AJ505411; Elsholtzia stauntonii Benth., U78690,

AJ505406; Lamium purpureum L., U78694,

AJ609175; Plectranthus barbatus Andrews,

U78698, AJ505378.

Martyniaceae: Craniolaria annua L., NEW, NEW; Ibicella lutea (Lindl.) Van Eselt., NEW, NEW;

Martynia annua L., NEW, NEW; Proboscidea

sabulosa Correll, NEW, NEW.

Orobanchaceae: Agalinis peduncularis (Benth.) Pennell, *, NEW; Agalinis tenuiflora Raf,

AY563927, *.

Paulowniaceae: Paulownia tomentosa (Thunb.) Steudl.,L36406,AJ431051.

Pedaliaceae: Ceratotheca triloba E. Mey. ex

Bernh., AY919281, AF482534; Harpagophytum

procubens DC. ex Meisn., NEW, NEW; Sesamum

indicum L., L36413, AJ431052; Sesamum radiatum

Schumach. & Thonn., NEW, NEW.

Phrymaceae: Diplacus aurantiacus Jeps., AF188186, AJ609163; Hemichaena fruticosa Benth., AJ617595, AJ609179; Lancea tibetica

Hook. f. & Thomson, AJ617599, AJ609174; Mazus

stachydifolius Maxim., AJ619559, AJ609167;

Phiyma leptostachya L., AJ617586, AJ431053.

Schlegeliaceae: Schlegelia parviflora (Oerst.) Monach., L36410, AJ431057.



110 Preliminary Chloroplast DNA Studies in Martyniaceae + Gutierriez

Scrophulariaceae: Leucophvllum frutescens

(Berland.) I. M. Johnst, AF123685, AJ609172.

Verbenaceae: Lantana c?mara L., *, AF225294; Lantana h?rrida H. B. & K., AF130152, *;

Stachytarpheta dichotoma (Ruiz & Pav?n) Vahl,

L36414, AJ299259; Verbena r?gida Spreng., AJ431065,AJ431065.



preliminary chloroplast dna studies in the flowering plant family martyniaceae (order lamiales)

Documents