Crisp & Chandler, Paraphyletic species 
821 
Table 3. Data matrix used in the cladistic analysis of Daviesia ulicifolia. Values for each character 
are standardised integers over the range 0 to 30 using the method of Thiele (1994). A polymorphism 
is indicated by 'p' (states 1 and 2). 
D. wyattiana 
30 
20 
? 
4 
29 
1 
30 
0 
30 
1 
1 
1 
D. acicularis 
12 
13 
17 
6 
11 
0 
0 
0 
5 
0 
1 
1 
D. arenaria 
2 
30 
16 
0 
12 
0 
0 
0 
8 
2 
0 
0 
D. arthropoda 
14 
26 
0 
2 
0 
0 
14 
0 
17 
0 
0 
0 
D. microcarpa 
6 
0 
30 
30 
8 
0 
0 
0 
10 
? 
1 
1 
D. uiicifolia: 
angustifolia 
6 
1 
25 
12 
11 
0 
0 
0 
0 
0 
0 
0 
desert 
7 
13 
22 
10 
5 
2 
7 
30 
8 
0 
0 
0 
grampians 
7 
20 
16 
5 
30 
1 
6 
0 
9 
0 
0 
0 
lofty 
9 
14 
20 
7 
27 
P 
7 
2 
8 
0 
0 
0 
NVP 
0 
20 
26 
3 
3 
0 
0 
0 
2 
0 
0 
0 
pilliga 
6 
26 
17 
1 
6 
0 
0 
0 
3 
0 
0 
0 
ruscifolia 
3 
18 
28 
4 
26 
0 
0 
0 
7 
0 
0 
0 
subumbellate 
6 
13 
24 
7 
22 
1 
5 
0 
5 
0 
0 
0 
typical 
5 
13 
22 
5 
16 
0 
3 
0 
6 
0 
0 
0 
yorke 
12 
28 
3 
2 
22 
1 
11 
0 
12 
0 
0 
0 
(de Queiroz & Donoghue 1988; Crisp & Weston 1993). Nevertheless one clade with 
a moderate bootstrap value of 67 included all forms of D. ulicifolia and the three 
autapomorphic species D. acicularis, D. arenaria and D. microcarpa (Fig. 2). Thus 
D. ulicifolia appears to be paraphyletic by exclusion of the latter species. Only the 
'Yorke' form is excluded from this clade, and it shows a sister-group relationship to 
D. arthropoda. As the species included within the D. ulicifolia clade are well separated, 
no re-rooting can make D. ulicifolia appear monophyletic, even if the distinct 'Yorke' 
is excluded from consideration. We tried constraining monophyly of D. ulicifolia, but 
this increased tree-length very substantially (68 extra steps), a significant difference 
which was not achieved in 100 randomised data sets (T—FTP < 0.01: Faith 1991). 
Other manipulations, such as selectively excluding combinations of species, did not 
substantially alter the relationships of the forms nor alter the paraphyly of D. ulicifolia. 
An alternative binary encoded data set (with fewer characters) produced a very 
unresolved tree but again showed D. ulicifolia as paraphyletic. 
Banksia integrifolia 
Thiele (1993b) and Thiele & Ladiges (1994) made a morphological analysis of the 
Banksia integrifolia (Proteaceae: Banksieae) complex, which broadly overlaps with 
D. ulicifolia east of the Great Dividing Range. Tlieir methods were essentially similar to 
those described above for D. idicifolia, using either binary or morphometric characters 
of adult leaves, fruits and juvenile leaves. Four phenetic clusters were recognised as 
taxa. Three of these overlapped in distribution and morphology. Only the most northern 
entity {'aquilonia') was found to be both morphologically and geographically disjunct, 
with no intermediates. The authors would have liked to segregate this as a species, 
but demurred on the basis of a cladistic analysis, which nested aquilonia deep within 
the complex (Fig. 3). Thus, B. integrifolia would have been rendered paraphyletic by 
