Introduced versus native subspecies of Codium fragile: how distinctive is the invasive subspecies tomentosoides?

After its introduction, the green alga Codium fragile (Sur.) Hariot ssp. tomentosoides (van Goor) Silva has spread widely on several temperate-zone, rocky shores where non-weedy conspecific subspecies occur (N.E. Atlantic, N.E. Pacific, S. Pacific). To determine how phenologically and morphologically distinctive the invasive alga was relative to native subspecies, I compared marine intertidal populations of C. fragile ssp. tomentosoides and the native C. fragile ssp. novaezelandiae (J. Ag.) Silva (hereafter referred to as ssp. tomentosoides and ssp. novae-zelandiae respectively on New Zealand shores in 1992, 1993 and 1995. On the North Island, the invasive ssp. tomentosoides is sparsely distributed on low intertidal benches on wave-protected shores in the Hauraki Gulf (east coast) in spring and summer, and thalli die back to the perennial holdfast in autumn. In contrast, the native ssp. novaezelandiae forms dense beds within the low intertidal mussel zone on wave-swept shores of Maori Bay (west coast), and fronds are perennial. Whereas ssp. tomentosoides has only a few fronds arising from the spongy basal hold-fast, ssp. novae-zelandiae thalli are composed of many fronds. The ssp. tomentosoides from the Hauraki Gulf is significantly more branched than comparably sized native conspecifics from Maori Bay. These phenological and morphological differences were used to predict the subspecific identity of C. fragile from three other locations on the North Island, two locations on the South Island, and four locations on S.E. Australian shores; microscopic examination of utricles was used to check the predictions. Seasonality and number of fronds per thallus are the most reliable characters for field identification of native vs invasive subspecies: perennial intertidal thalli with large numbers of fronds are indicative of native subspecies for different geographic regions.