WOOD ANATOMY AND STEM OF CHLORANTHUS - SUMMARY OF WOOD ANATOMY OF CHLORANTHACEAE, WITH COMMENTS ON RELATIONSHIPS, VESSELLESSNESS, AND THE ORIGIN OF MONOCOTYLEDONS

In contrast to the monopodial Ascarina and Hedyosmum, Chloranthus and Sarcandra are sympodial. Sarcandra and C. erectus have woody canes of finite duration, whereas other species of Chloranthus have shoots of one year's duration; these latter species have second year wood only on rhizomes, not on upright shoots. Rhizome portions transitional to upright stems were selected for study. Chloranthus erectus has abundant septate fibre-tracheids, C. japonicus none, and two other species a few. Chloranthus (and Sarcandra) have rays of two distinct sizes in wood: rays that are extensions of primary rays, and uniseriate and biseriate rays in fascicular areas. Wood anatomy of each of the four genera can be characterised, and is summarised in the form of a key. Except for primitiveness of vessels, wood of Chloranthaceae is very similar to that of Lactoridaceae and Piperaceae, and this probably indicates a close phyletic relationship. The large rays of chloranthaceous wood, little modified from primary rays and with upright cells predominantly, are indicative of some degree of herbaceousness and some degree of secondary woodiness. Scattered bundles and multilacunar nodes, characteristics of monocotyledons, are absent in Chloranthaceae but present in Piperaceae. The sympodial habit of Chloranthus and Sarcandra, and the presence of vessels in roots but not in stems of Sarcandra are conditions like those basic to origin of monocotyledons. The possibility that Chloranthaceae are close to Piperales and that these groups are close to origin of monocotyledons should be considered. Some cladists have hypothesised that secondary vessellessness is polyphyletic in dicotyledons. While these cases are theoretically possible, the histological and ecological scenarios that must be hypothesised for these events are ignored by cladists; most of these scenarios are unlikely for reasons explored here, although a few are still worthy of consideration. Stem endodermis is reported for three species of Chloranthus.