N. Wikstrom et P. Kenrick, Relationships of Lycopodium and Lycopodiella based on combined plastid rbcL gene and trnL intron sequence data, SYST BOT, 25(3), 2000, pp. 495-510
The relationships of Lycopodium and Lycopodiella (Lycopodiaceae) were inves
tigated based on two plastid data sets (rbcL gene and trnL intron) from a r
epresentative sample of 21 species. Separate and combined analyses of the d
ata reveal consistent patterns of relationship. There is strong support for
monophyly of Lycopodium and Lycopodiella. There is also support for monoph
yly of species groups or sections sensu Ollgaard (Lycopodium, Diphasium, Ma
gellanica, Complanata, Lycopodiella, and Campylostachys). The combined data
provide new evidence of relationships between subgeneric groups. In Lycopo
dium, section Pseudodiphasium groups with section Magellanica section Obscu
ra groups with section Diphasium, and section Annotina groups with section
Lycopodium. in Lycopodiella, sections Lateristachys and Caroliniana group w
ith section Campylostachys and this group is sister to section Lycopodiella
. Tentative calibration of the phylogenetic tree using fossil evidence indi
cates a minimum age of Early Jurassic (208 Myr) for the split between Lycop
odium and Lycopodiella. Reticulate fossil spores from Upper Permian records
are potentially of Lycopodium affinity and indicate that early cladogenesi
s in Lycopodium may be even older. An evaluation of biogeographic and phylo
genetic patterns in these two genera shows a striking difference from that
in Huperzia. Sections within Lycopodium and Lycopodiella have broad geograp
hic distributions, whereas molecular data partition the much larger Huperzi
a group into predominantly neotropical and paleotropical clades.