Basal Angiosperms-Magnoliids

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Department of Plant Biology and Biodiveriuty
Management,College of Natural Sciences,
AAUEthioipia

•  
• Phylogeny of Angisperms (PBBM 802)
• Asnake Tsegaw (asnake.tsegaw_at_aau.edu.et)

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Content

• What is the APG classification?
• Basal angiosperms
• ANITA (basal families)
• Magnoliid complex

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APG II Modern Plant Taxonomy

• This is a revision of plant taxonomy basically
  avoiding the nomenclature of taxonomy and working
  primarily at the level of order and family. It
  stresses cladistic trees rather than levels of
  taxonomy.
• Published in 2003
• Made clarifications from the 1998 paper
• Basically just incorporated newer molecular data
• Always changing and being revised, but with a
  good standard, that of the monophyletic group

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An update of the Angiosperm Phylogeny Group
classification for the orders and families of
flowering plants APG IITHE ANGIOSPERM PHYLOGENY
GROUP
APG II Modern Plant Taxonomy
A revised and updated classification for the
families of the flowering plants is provided.
Newly adopted orders include Austrobaileyales,
Canellales, Gunnerales, Crossosomatales and
Celastrales. Pertinent literature published since
the first APG classification is included, such
that many additional families are now placed in
the phylogenetic scheme. Among these are
Hydnoraceae (Piperales), Nartheciaceae
(Dioscoreales), Corsiaceae (Liliales),
Triuridaceae (Pandanales), Hanguanaceae
(Commelinales), Bromeliacae, Mayacaceae and
Rapateaceae (all Poales),
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What are basal angiosperms?

• Basal angiosperms
• This is a polyphyletic group of families that are
  ancestral to all other angiosperms
• Those groups that are now shown to be the oldest
  diverging paraphyletic!
• Are less derived and include the flowering plant
  belonging to the oldest lineage
• Example Magnolia primitive

• polypetaly
• hypogyny
• Actinomorphic

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What are basal angiosperms?

• Exhibit a suite of primitive character states
• Many parts at each whorl
• Separate, unsealed carpels
• Follicle fruits

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What are basal angiosperms?

• Laminar stamens
• Tracheids, no vessel elements
• Pollen grains single pored, apertured,
• furrowed not triaperturate, tricolpate

Pollen of Horse chestnut
A Grass Pollen
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ANITA Basal Angiosperms

• The ANITA group forms a basal grade
  (paraphyletic)
• Name derived from initials of members

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Number of families, genera, species
ANITA Basal Angiosperms
Amborellales 1/1/1 Nymphaeales 3/7/74 Austrobailey
ales 3/5/100 1/4/75

• Amborellales
• Amborellaceae 1 g, 1 spp
• Nymphaeales
• Hydatellaceae 2 g, 10 spp
• Cabombaceae 2 g, 6 spp
• Nymphaeaceae 3 g, 58 spp
• Austrobaileyales
• Austrobaileyaceae 1 g, 2 spp
• Schisandraceae 3 g, 92 spp
• Trimeniaceae 2 g, 6 spp

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ANITA Basal Angiosperms
First branch - Amborella

• As the sister to all other extant angiosperm
• Darwins famously called the origin of
  angiosperm as Abominable mystery
• Even describe as the Holy Grail of botany
• Different from most Laurales in spiral arranged
  flora
• While fused tissues in most angiosperm
• ( plesiomorphy for the angiosperm)
• Features which are synapomorphy for the extant
  angiosperm except Amborella
• Vessel
• Pollen grain with reticulate tectum and
• Ethere oil cells ( also for Nymphaeceae)

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ANITA Basal Angiosperms
First branch - Amborella

• Nearly all multi-gene analyses ( slow-
  evolving gene, protein coding, rRNA, also-
  rapid-evolving plastid gene, non-coding) of
  basal angiosperms have identified Amborella as
  the sister to all other extant of Angiopsrms with
  varying level of support)
• Example egg formation in the angiosperm
  Amborella resembles that of the gymnosperm

• One speices, Amborella trichopoda
• Endemic to New Caledonia
• Functionally dioecious
• Fruit an aggregate of drupes
• Wood lacks vessels
• Apocarpic, not sealed
• perianth spiralled

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ANITA Basal Angiosperms
Second branch - Nymphaeales
Nymphaea odorata

• Flowers bisexual, radial, with a long pedicel
• Petals usually 8 to numerous
• Stamens 3 to numerous
• Seeds usually operculate (opening by a cap)
• Hairs simple usually producing mucilage (slime)
• floating or submersed leaves
• air cavities in tissue
• lack of vessels

• Includes ornamental water lily
• Widespread distribution except arid regions
• Aquatic, rhizomatous herbs

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ANITA Basal Angiosperms
Austrobaileyales (100 species) Illiciaceae

• Aromatic Trees or shrubs
• Source of star anise ( cooking and medicine,
  source of oil )
• Although the traditional Illiciaceae and
  Schisandraceae have typically been united in
  Illiciales, a relation between these taxa and
  Austrobaileya and Trimenian had not been
  suspected
• No morphological synapomorphies have been
  identified for this clade, despite the strong
  molecular support for its monophyletic relaton.

• Flowers Inflorescences only 1-3 flowers, with
  tepals
• Filaments poorly differentiated from the anthers
• Carples 7 to numerous, not sealed
• Fruit a star like aggregate of 1-seeded follicles

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Phylogeny of Angiosperms
Basal angiosperms
Fusion
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Modern angiosperm relationships
Basal Lineages
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Eudicots
Monocots
1
2
3

1. Amborellaceae
2. Nymphaeaceae
3. Austrobaileyales
4. Magnoliid complex

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Magnoliid Basal Angiosperms

• Magnoliid
• The clade comprise most of lineage referred as
  primitive angiosperm
• Previously ( before phylogentics)
• Are loosly associated
• Relation among the families and order were not
  clear
• Contained group that are not part of magnoliid
  clade
• Compositions and interrelationship of magnollid
  were not clear
• After data set of at least five gene for broad
  sample of taxa, (i.e. phylogentics)
• Age of the clade and high level of extinction
  make the reconstruction and recognition of the
  clade more difficult
• Within Magnoliid, Magnoliales and Laurales, and
  Piperales and Canellales are sister

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Magnoliid Basal Angiosperms

• Magnoliid
• Magnollids share some traits with basal
  angiosperm but are more close related to monocot
  and eudicot
• Example carpel are free from one another unlike
  most contemporary angiosperm
• Abominable mystery- two main competing
  hypotheses have take center stage
• Woody magnoliid hypotheses (Modern spp.
  reprsentative Mgnolia grandiflora, Liriodendron
  tulipifera, Anna crassiflora) agnaist Paleoherb
  hypotheses ( Piper nigrum, Ausarum caudatum,
  Aristolochia arborea)

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Magnoliid Basal Angiosperms

• Magnoliales
• Trees or shrubs, large showy flowers
• 6 families
• Laurales
• Trees or shrubs, smaller bisexual or unisexual
  flowers
• 7 families
• Canellales
• Trees or shrubs, flowers 5-merous to many
• 2 families
• Piperales
• Herbs or small shrubs, reduced often 3-merous
  flowers
• 5 families

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Magnoliids Canellales Canellaceae 5 g, 13
spp Winteraceae 5 g, 80 spp Laurales Calycanthacea
e 5 g, 11 spp Siparunaceae 2 g, 75
spp Gomortegaceae 1 g, 1 spp Atherospermataceae 7
g, 16 spp Monimiaceae 22 g, 200 spp Hernandiaceae
5 g, 55 spp Lauraceae 50 g, 2500
spp Magnoliales Myristicaceae 20 g, 475
spp Magnoliaceae 2 g, 227 spp Degeneriaceae 1 g,
2 spp Himantandraceae 1 g, 2 spp Eupomatiaceae 1
g, 3 spp Annonaceae 129 g, 2220
spp Piperales Hydnoraceae 2 g, 7
spp Aristolochiaceae 6 g, 480 spp Piperaceae 5 g,
3600 spp Saururaceae 4 g, 6 spp
Number of families, genera, species
Canellales 2/10/93 Laurales 7/91/2858 Magnoliales
6/154/2929 Piperales 4/17/4090
Magnoliids
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Magnoliid Basal Angiosperms

• Magnoliaceae (Magnoliales)
• Remainder of basal angiosperms
• Flower parts often in 3s or many
• One pollen aperture
• Two cotyledons
• Tropical and temperate
• Trees or shrubs

• aromatic - ethereal oils, (ranalian smell)
• Flowers born spirally on an elongate axis
• beetle or fly pollination common

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Magnoliid Basal Angiosperms
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Magnoliaceae (Magnoliales)
Magnoliid Basal Angiosperms

• Flowers born spirally on an elongate axis
• Widely distributed in temperate and tropical
• One of the oldest
• Its feature thought to have been present in the
  ancient angiosperm
• So, it is placed at the basal of angiosperm

• The delimitation of genera in Magnoliaceae has
  changed, based on molecular studies, to the
  recognition of just two genera, Magnolia (225
  species) and Liriodendron (2 species).

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Magnoliaceae (Magnoliales)
Magnoliid Basal Angiosperms
g
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• Characteristics Leaves usually alternate,
• entire with stipules
• perianth tepals spirally arranged
• Flowers bisexual, radial, solitary, distinct
• 9-15 tepals in whorls of 3
• androecium o a f laminar stamens
• Carpels usually numerous, on elongate receptacle
• fruits of one flower cone or aggregate of
  follicles
• Two genera Magnolia and Liriodendron

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Magnoliid Basal Angiosperms
Magnoliaceae (Magnoliales)
Magnolia grandiflora Note many flower parts on
an enlarged receptacle, leaf-like stamens.
Magnolia grandiflora
Within the family, Magnolia is diagnosed
by -presence of a red or orange fleshy seed
coat -follicles opening along the abaxial seam
Magnolia grandiflora Young fruits (follicles)
developing. Stigmas are turning brown, anthers
are drying up.
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Magnoliid Basal Angiosperms
Magnoliaceae (Magnoliales)
Liriodendron tulipifera, Tulip Tree, Note
tulip-shaped leaf, radial symmetry, woody, many
flower parts, yellow.
Cross section Note expanded receptacle, many
stamens and carpels, leaf-like stamens, radial
symmetry.
Carpels separate at maturity producing samaras
fruit.
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Magnoliid Basal AngiospermsAnnonaceae
(Magnoliales) - custard apples

• About 120 genera and over 1000species
• trees Or Shrubs
• Confined to tropical and sub-tropical areas
• characteristic of lowland evergreen forests in
  Asia and Africa
• large, woody pantropical family
• Perianth in 3 sets of three tepals

Annona cherimola
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Magnoliid Basal AngiospermsAnnonaceae
(Magnoliales) - custard apples

• Characteristics Simple Alternate leaves
• Flowers actinomorphic hypogynous
• Numerous stamens, laminar
• spirally arranged on the receptacle
• Carpel free, not fusing into fleshy syncarp at
  maturity
• An aggregate fruit of berries or a syncarp
  formed by connation of carpels, aromatic
• Aromatic nodding flowers and numerous floral
  parts.
• Genera and species Annona (120 species),
  Artabotrys, Uvaria (175 species) and Xylopia (150
  species).

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Magnoliid Basal Angiosperms

• Myristicaceae (Magnoliales) - nutmeg
• tropical family with members in Central America,
  the northern half of South America, Central
  Africa, Asia
• The largest genus, Myristica (including M.
  fragrans, the source of nutmeg and mace), has
  about 175 species.
• Myristica fragrans - nutmeg, mace
• mace from aril
• nutmeg from seed

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Magnoliid Basal Angiosperms Degeneriaceae
(Magnoliales)

• Degeneriaceae consists of one genus in Fiji.
  Degeneria vitiensis, as the species name
  indicates, was found on Vtit Levu, the largest
  island of the Fijian archipelago.
• It is a relatively common tree and used for
  timber.
• A second species, D. roseiflora, was described in
  1988 on different Fijian islandsnamely, Vanua
  Levu and Taveuni.
• fairly common timber tree
• differs from the first species in
• having magenta or pink flowers,
• smaller fruits, and bark of a
• different colour.

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Magnoliid Basal Angiosperms
Eupomatiaceae and Himantandraceae
(Magnoliales)

• Eupomatiaceae consists of one genus, Eupomati,
  with two species.
• Eupomati laurina a common rainforest shrub in New
  Guinea and Australia.
• Eupomati benneti, is much less common and is
  restricted to Australia
• Himantandraceae contains one genus, Galbulimima
  (also known as Himantandra), found in the Molucca
  Islands of Indonesia, Malaysia, New Guinea, and
  northeastern Australia.

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Magnoliid Basal Angiosperms Canellales
(Canellaceae and Winteraceae )

• Canellales
•  formerly called Winterales, 
• relatively little economic importance.
• Winteraceae
• have little economic importance.
• The leaves peppery taste, discourages browsing
  animals.
• The leaves and bark of the West Indian Canella
  alba (wild cinnamon), use as a condiment and for
  medicinal purposes.
• Drimys winteri variety chilensis is cultivated in
  many parts of the world in gardens and
  arboretums.
• A small bushy tree in cultivation, and has
  attractive white-petaled flowers

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Magnoliid Basal AngiospermsLaurales

• Members of Laurales are characterized by
• woodiness,
• aromatic parts, and
• a single strand of conducting tissues continuing
  from the stem into the leaf.
• Lauraceae and Monimiaceae together constitute
  most of the genera in this order.

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Magnoliid Basal AngiospermsLauraceae (Laurales)

• 50 genera and 2500 species
• Widespread in tropics and subtropics
• Trees or shrubs, or twining parasitic vine
• Source of many spices
• bay leaves, cinnamon, camphor, sassafras
• fruits avocado
• Cassytha, a rootless vine like stem parasite with
  vestigial scale like leaves.
• L. nobilis (sweet bay tree, or bay laurel), a
  native of the Mediterranean
• Sassafras, a genera, and its oil was used
  medicinally, and a tea from the bark and twigs.

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Magnoliid Basal AngiospermsLauraceae (Laurales)

• Stamens with basal nectar appendages
• Fruit 1 seeded berry or drupe and valved anthers
• Leaves alternate and spiral, entire, without
  stipules
• and aromatic (ethereal oils)
• Flowers concave receptacle that often enlarges
  in fruit anthers dehiscing via pores with flaps
  sticky pollen pollen without apertures carpel
  1 embryo large, endosperm lacking.
• Significant features ethereal oils
• Special uses Cinnamon (Cinnamomum) and bay
  (Laurus nobilis)
• used as spices avocado (Persea)

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Magnoliid Basal AngiospermsLauraceae (Laurales)
Cinnamon (Cinnamomum zeylanicum), bushy evergreen
tree ,native to Sri Lanka (Ceylon), for the spice
consisting of its dried inner bark. The spice is
light brown in colour and has a delicately
fragrant aroma and warm, sweet flavour

• Persea americana,
• These flowers are dichogamous
• a tree native to the Western Hemisphere from
  Mexico south to the Andean regions.
• have greenish or yellowish flesh with a buttery
  consistency and a rich, nutty flavour.
• They are often eaten in salads, and in many parts
  of the world they are eaten as a dessert.

Parasitic climber , with leafless filiform stem
Cassytha filiformis
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Magnoliid Basal Angiosperms Monimiaceae
(Laurales)

• The second largest family Monimiaceae has 22
  genera and 200 species,
• less than 10 percent of Laurales species.
• also found in tropical and subtropical regions,
• it is less extensively distributed
• occurs mainly in the warmer areas of the Southern
  Hemisphere.
• The type genus, Monimia, is restricted to the
  Mascarene Islands

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Magnoliid Basal Angiosperms Siparunaceae
(Laurales)

• 75 species in two genera.
• Glossocalyx, from tropical West Africa, has four
  species.
• the genus Siparuna, found in Mexico, Central
  America, and tropical South America.
• are trees or woody vines
• opposite, mostly serrate leaves.
• The flowers are unisexual
• stamens glands are not present at the base of
  the
• The hypanthium becomes woody and splits.

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Magnoliid Basal Angiosperms Hernandiaceae and
Atherospermataceae (Laurales)

• The remaining four families have a combined total
  of 83 species (Hernandiaceae, Atherospermataceae,
  Calycanthaceae and Gomortegaceae)
• Hernandiaceae (55 species) is a pantropical
  family of trees, shrubs, and some lianas.
• The largest genus, Hernandia (22 species), is
  distributed in Central and South America, the
  West Indies, West Africa, Indo-Malaysia
• Atherospermataceae includes 6 or 7 genera and 16
  species, which are native to Australia, New
  Guinea, New Zealand, New Caledonia, and Chile.

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Magnoliid Basal Angiosperms Calycanthaceae
(Laurales)

• includes 75 species in two genera.
• the family, has a discontinuous distribution
• Calycanthus (strawberry shrub, or sweet shrub,)
  is found in United States,
• Chimonanthus and Sinocalycanthus occur in China.
  
• Glossocalyx, from tropical West Africa, has four
  species.
• the genus Siparuna, found in Mexico, Central
  America, and tropical South America.
• Deciduous shrubs
• Leaves opposite without stipules
• Flowers bisexual with hypanthium
• 15-30 tepals, spiral
• Grown ornamentally

Calycanthus occidentalis
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Magnoliid Basal Angiosperms Gomortegaceae
(Laurales)

• The single species of Idiospermum is a very rare
  evergreen species from Queensland, Austl.
• Gomortegaceae, or the queule family, consists of
  a single species, Gomortega keule, which is a
  rare species native to central Chile.

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Magnoliid Basal AngiospermsAristolochiaceae
(Piperales) - birthwort, wild ginger

• Characteristics aristolochic acids
• (bitter, yellow, nitrogenous compounds)
• Leaves alternate and spiral, sometimes lobed,
  palmate
• Flowers Highly modified, showy, fused sepals,
  radial or bilateral, tubular, and S-shaped or
  pipe shaped calyx tube corolla usually lacking
  or vestigial ovary inferior or half-inferior, of
  3-6 connate carpels ovules numerous filaments
  more or less adnate to style.
• Significant features dead meat carrion
  coloration attracts insects, usually flies.
  Trap-flowers.
• Special uses ornamentals some medicinal uses.
• Fruit a capsule

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Magnoliid Basal AngiospermsAristolochiaceae
(Piperales) - birthwort, wild ginger
(Dutchmans Pipe or Birthwort Family)

• Asarum (wild ginger)
• stemless perennial with aromatic rhizomes
• flowers actinomorphic
• filament tips extending beyond the anthers
• ovary inferior
• fruit a fleshy capsule, seeds large
• Characters of Aristolochia
• Tropical or warm temperate regions
• Perennial herbs or shrubs, twining or climbing
• Calyx tubular, greenish or purplish
• Anthers sessile, strongly adnate to the short and
  fleshy style
• trap flowers

Asarum and Saruma are herbaceous genera of the
Northern Temperate Zone, most diverse in eastern
Asia.
Saururus cernuus
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Aristolochiaceae
Fly pollination - stigma and flower withers and
flies can leave
Branch 2-3 long with slender pendulous flowering
stem, ornamentals
Aristolochia
fragmentation of rhizomes in the lizards-tail
family assures vegetative propagation
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Magnoliid Basal AngiospermsSaururaceae
(Piperales) - lizards-tail

• native to North America and Southeast Asia.
• It includes five genera and six species,
• most of them aromatic herbs with creeping
  rhizomes
• the plants generally inhabit wet areas.
• Perennial herbs of north temperate
• In Saururus the elongate cluster bends at its tip
  to suggest the fanciful appellation lizards
  tail.
• Cordate leaves
• Reduced flowers on showy inflorescence

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Saururaceae and Piperaceae
Magnoliid Basal AngiospermsSaururaceae
(Piperales) - lizards-tail

• Saururaceae and Piperaceae are closely related.
• Their inflorescence are slender, spike like, and
  covered with inconspicuous, closely appressed
  flowers.
• the floral spikes are showy,
• in several Saururaceae species the spikes
  resemble a single flower because of the expanded
  bracts that arise below the inflorescence.
• Other features common to most species of
• these two families include
• a sheathing leaf base,
• lack of sepals and petals, bisexual flowers,
• and presence of one erect ovule per ovary
  chamber.

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Magnoliid Basal AngiospermsPiperaceae
(Piperales) - pepper, pepperomia

• 6 genera and 2020 species
• most of them in the large genera Pepperomia and
  Piper (the black pepper genus)
• Widely distributed in tropics and subtropics
• Herbs to small trees, or epiphytic
• Reproduction by seeds is the main method of
  species dispersal in Piperales
• Black, white pepper and betel nut source

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Magnoliid Basal Angiosperms Hydnoraceae
(Piperales) - lizards-tail

• Small family with seven species in two genera.
• They are terrestrial parasitic plants that lack
  leaves and chlorophyll.
• The large flowers have a single three-parted
  perianth whorl and an inferior ovary
• They are foul smelling and are pollinated by
  flies and beetles.
• Prosopanche occurs in Central and South America
• Hydnora occurs in Africa, Madagascar, and the
  Arabian Peninsula.

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Piperaceae
Inflorescence a spike densely covered with minute
flowers
Leaves alternate, pinnate to palmate venation.
Nodes Swollen and Jointed
Each flower with a peltate bract.
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Magnoliid Basal AngiospermsPiperaceae
(Piperales) - pepper, pepperomia

• charcterstics vascular bundles of more than 1
  ring, Inflorescences densely covered in minute
  flowers
• Leaves usually alternate and spiral, entire
• Flowers minute, densely packed in a thick spike
  or spadix no perianth (!) carpels 1-4, connate,
  with 1 ovule per gynoecium, basal placentation
  fruit usually a drupe.
• Significant features stem with bundles in gt1
  ring or scattered
• Special uses Piper nigrum (black pepper) Piper
  betle (betel pepper), ornamentals (Peperomia)
• Fruit a drupe

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Magnoliid Basal AngiospermsPiperaceae
(Piperales) - pepper, pepperomia
Peperomia and Piper Importance Various species
of Piper are sources of spice.

• Piper nigrum, Black pepper, Piperaceae
• Erect shrub, climber, leaves up to at least 11cm
  long
• Aromatic when crushed
• Stipule present, when lost it leave scars
• Piper longum, the dry spike used as one
  ingredient in spicing tea

• Peperomia
• not aromatic
• Stipule absent
• Leaves not more
• than 8cm

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END OF THE BASAL LINEAGS
Basal Lineages
Monocots
Eudicots