Editing Microphylls and megaphylls (section)

===Evolution===
The '''enation theory''' of microphyll evolution posits that small outgrowths, or [[enation]]s, developed from the side of early stems (such as those found in the [[Zosterophyll]]s).<ref name=Stewart&Rothwell>WN Stewart & GW Rothwell (1993) Palaeobotany and the evolution of plants. 2nd edition. Cambridge University Press.</ref>  Outgrowths of the protostele (the central vasculature) later emerged towards the enations (as in ''[[Asteroxylon]]''),<ref name=Stewart&Rothwell/> and eventually continued to grow fully into the leaf to form the mid-vein (such as in ''[[Baragwanathia]]''<ref name=Stewart&Rothwell/>).<ref name=Kaplan2001/>  The fossil record appears to display these traits in this order,<ref name=Stewart&Rothwell/> but this may be a coincidence, as the record is incomplete. The '''telome theory''' proposes instead that both microphylls and megaphylls originated by the reduction; microphylls by reduction of a single ''telome'' branch, and megaphylls by evolution from branched portions of a telome.<ref name=Stewart&Rothwell/>

The simplistic evolutionary models, however, do not correspond well to evolutionary relationships. Some genera of ferns display complex leaves that are attached to the pseudostele by an outgrowth of the vascular bundle, leaving no leaf gap.<ref name=Kaplan2001/> Horsetails (''[[Equisetum]]'') bear only a single vein, and appear to be microphyllous; however, the fossil record suggests that their forebears had leaves with complex venation, and their current state is a result of secondary simplification.<ref name=Taylor1993>{{cite journal
 | author = Taylor, T.N.
 |author2=Taylor, E.L.
 | year = 1993
 | title = The biology and evolution of fossil plants
}}</ref>  Some [[gymnosperm]]s bear needles with only one vein, but these evolved later from plants with complex leaves.<ref name=Kaplan2001/>

An interesting case is that of ''[[Psilotum]]'', which has a (simple) protostele, and enations devoid of vascular tissue.  Some species of ''Psilotum'' have a single vascular trace that terminates at the base of the enations.<ref name=Gifford1989/> Consequently, ''Psilotum'' was long thought to be a "[[living fossil]]" closely related to early land plants ([[rhyniophyte]]s). However, genetic analysis has shown ''Psilotum'' to be a reduced fern.<ref name=Qiu1999>{{cite journal
 | author = Qiu, Y.L.
 |author2=Palmer, J.D.
 | year = 1999
 | title = Phylogeny of early land plants: insights from genes and genomes
 | journal = Trends in Plant Science
 | volume = 4
 | issue = 1
 | pages = 26–30
 | doi = 10.1016/S1360-1385(98)01361-2
 | pmid = 10234267
}}</ref>

It is not clear whether leaf gaps are a [[Homology (biology)|homologous trait]] of megaphyllous organisms or have evolved more than once.<ref name=Kaplan2001/>

While the simple definitions (microphylls: one vein, macrophylls: more than one) can still be used in modern botany, the evolutionary history is harder to decipher.
<gallery>
File:Vein sceleton hydrangea ies.jpg|Megaphylls have a complex network of veins.
File:Psilotum.jpg|''[[Psilotum]]'' has secondarily lost leaves, and bears [[enation]]s resembling the microphylls of early land plants.
</gallery>