The Pacific Northwest tall shrub Acer circinatum (vine maple) can host diverse and abundant epiphyte communities. A chronosequence approach revealed that these communities gradually shift in composition as the shrub progresses through its life cycle. Different epiphytic life forms occupy different spatial and temporal niches on shrub stems. These life forms generally shift upwards along the shrub stem as the stem ages and develops, in accordance with the similar gradient hypothesis. We postulate the following sequence of events. An initial wave of colonization occurs as new substrate is laid down. Over time, superior competitors gradually engulf and overgrow competitively inferior primary colonizers. Concurrently, shrub stem microclimate changes as shrub stems grow, age, and layer, causing the processes of competition and colonization to shift in favor of different epiphytic life forms during different life stages of the shrub stem. We define four separate shrub stem life stages: life classes 1–4 describe, respectively, young upright ‘whips’; vigorous, upright, mature stems; declining stems beginning to bend towards the forest floor; and horizontal, decadent stems. As space on the shrub stem is filled through growth and colonization, interspecific competition intensifies. Successful competitors persist and spread, while poor competitors are increasingly restricted to the stem tips, where interspecific competition is less intense. In these forests, Usnea, green-algal foliose lichens, and moss tufts excel as the primary colonizers and become common on the outer portions of shrub stems over time, as long as the overstory is not too dense. Moss mats are also good primary colonizers, but excel as secondary colonizers, often coming to dominate decadent shrub stems. Although all life forms can be primary colonizers, the remaining forms (cyanolichens, liverworts, and Antitrichia curtipendula) are effective secondary colonizers. Liverworts are also effective competitors, but less so than the moss mats on the most decadent stems. Cyanolichens appear to benefit from the aging and decline of shrub stems. The ability of vine maple to continually generate new young stems through basal sprouting and layering make it a varied and dynamic substrate for epiphytes. Such shrubs may act as epiphyte dispersal agents, with the potential to affect epiphyte continuity within forest stands that have experienced large disturbances.
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Vol. 104 • No. 2