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Speciation in the Australian Broadleaved

Paperbark Complex (Myrtaceae)

Bort (Robert) Edwards


PhD candidate, The University of Queensland, Brisbane

Dr Lyn Cook (UQ), Prof Mike Crisp (ANU), Mr Lyn Craven (CSIRO)
The Melaleuca leucadendra complex
The Melaleuca leucadendra complex
Nested within Melaleuca s.l.

Non-monophyletic

Crown age of approx 6MYA, coinciding with


aridification and expansion of dry sclerophyll vegetation

6 MYA
Melaleuca s.l.

(Edwards et al., 2010) (Cook et al., 2008)


Broadly sympatric

12 morphospecies overlap on Nth Cape York


Typically found in habitats subject to regular flooding,
local-scale segregation appears associated with landform

Edaphic influence: - water availability (Franklin et al, 2007)


Excellent group to study the role of niche differentiation
in speciation and/or the maintenance of species boundaries

Gene flow and structure across large geographic areas

The influence of landforms and geographical barriers to geneflow


Testing Species Limits and Models of Speciation:
Testing Species Limits and Models of Speciation:

Test hypothesis 2 using genetic data

Explore hypotheses 3 and 4 using


genetic data and ecological niche modeling

sampling across range to capture geographic variation/divergence


~200 individuals sampled from fresh and herbarium
material
-14 morphospecies
Number of samples roughly proportional to each species
range
One chloroplast region (psbA-trnH)
One nuclear region (ITS) and cpDNA (psbA-trnH)
cpDNA

Minimum spanning network, 14 species


cpDNA

Minimum spanning network, 14 species, cpDNA


cpDNA

Minimum spanning network, 14 species, cpDNA


cpDNA

Minimum spanning network, 14 species, cpDNA


ncDNA

Minimum spanning network, 14 species


Both chloroplast and nuclear DNA show strong signals of genetic
differentiation with moderate to low levels of incomplete lineage sorting

Reject hypothesis 2

Case study to investigate the possibility of ecological differentiation/speciation..


M. argentea and M. fluviatilis
Species Limits

cpDNA
Haplotypes shared between species almost exclusively
large internal (putatively ancestral)

Internal haplotypes are also geographically wide-spread

Fst = 0.145

Amova
among species: 14.48%
within species: 85.52%
(P=0.02)
Species Limits

ncDNA

Fst = 0.094

Amova
among species: 9.41%
within species: 90.59%
(P=0.018)

shared large widespread internal haplotypes,


private, localised external haplotypes

- Recent divergence
- Distinct species with no ongoing gene flow but incomplete lineage sorting
given that we are dealing with distinct species, with a region of overlap and no apparent physical barrier to gene flow:

Hypothesis 3 Hypothesis 4

there is contemporary difference there MAY (or may not) be


between species niches contemporary difference between
species niches

there is NO contemporary ecological there is NO contemporary ecological


barrier to gene flow (unsuitable barrier to gene flow (unsuitable
ribbon of habitat) ribbon of habitat)

there has been NO historical barrier there HAS been a historical barrier
to gene flow to gene flow with secondary contact
Comparing ecological niches

- niche values are estimated via MaxEnt (Phillips et al,


2006)

- observed niche values within and between species


ranges are compared against randomised distributions
generated via ENMtools (Glor & Warren, 2010)

null distribution

obs value

More different More similar


Is there a difference between the background environments from which each
species is drawn?

The background environment the species are drawn from


are no more different than expected by chance
Is there contemporary difference between the species’ specific niches?

Species’ niches are more different than expected by chance


Is there contemporary ecological barrier to gene flow?

Ribbon of overlap between species is no more or less


Different than the areas of species ranges either side
Hypothesis 3 Hypothesis 4

there is contemporary difference there MAY (or may not) be


between species niches contemporary difference between
species niches

there is NO contemporary ecological there is NO contemporary ecological


barrier to gene flow (unsuitable barrier to gene flow (unsuitable
ribbon of habitat) ribbon of habitat)

there has been NO historical barrier there HAS been a historical barrier
to gene flow to gene flow with secondary contact
PAST barrier to gene flow with secondary contact:
Species are linked to drainage systems

Populations diverge in allopatry across


the Great Dividing Range

Drainage capture with diversion of some


east-flowing headwaters to the west
allows secondary contact

As suggested for some species of fresh-water


Fish in the region (Hughes & Hillyer 2006;
Hurwood & Hughes, 1998)

Further insight needed to differentiate


processes
Morphological species limits within the Broadleaf Paperbark complex
(and other genera within Myrtaceae) are difficult to define:

- generalist morphologies
- generic (flexible and successful!) characters
- large/massive effective population sizes, intraspecific variation

However molecular data can be applied to species limits, as well as:

- ecological adaptation/specialisation
- phylogeography
- evolution in response to historical processes (climate, vicariance etc)
University of Queensland Postgraduate
Research Scholarship

Australian Systematic Botany Society’s


Hansjorg Eichler Research Fund

The Cook Lab (The University of Queensland)

Anna Kearns

The collectors who have contributed


material to various Australian herbariums

References
1 Cook L, Morris D, Edwards R, Crisp M (2008) Reticulate evolution in the natural range of the invasive
2 wetland tree species Melaleuca quinquenervia. Mol Phylogenet Evol 47: 506-522.
3
4 Franklin D, Brocklehurst P, Lynch D, Bowman D (2007) Niche differentiation and regeneration in the
5 seasonally flooded Melaleuca forests of northern Australia. J Trop Ecol 23:457 -467
6
7 Glor R, Warren D (2010) Testing Ecological Explanations for Biogeographic Boundaries. Evolution 65:673 -
8 683.
9
10 Hughes J, Hillyer M (2006) Mitochondrial DNA and allozymes reveal high dispersal abilities and historical
11 movement across drainage boundaries in two species of freshwat er fishes from inland rivers in
12 Queensland, Australia. J Fish Biol 68:270 -291.
13
14 Hurwood D, Hughes J (1998) Phylogeograph y of the freshwater fish, Mogurnda adspersa, in streams of
15 northeas tern Queen sland, Australia: evidence for altered dra inage pattern s. Mol Ecol 7:1507 -1517.
16
17 Phillips S, Anderson R, Schapire R (2006) Maximum entropy modeling of species geographic distributions.
18 Ecol Model 190:231-259.
19
20 Edwards R, Craven L, Crisp M, Cook L (2010) Melaleuc a revisited: cpDNA and morph ological data co nfirm
21 that Melaleuca L. (Myrtaceae) is not monophyletic. Taxon 59:744-754

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