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Germination Depth Vs. Temperature Requirements in Naturally Dispersed Seeds of Leucospermum Cordifolium and L

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Germination Depth Vs. Temperature Requirements in Naturally Dispersed Seeds of Leucospermum Cordifolium and L S. Afr. J. Bot. , 1987 , 53(2): 119-124 119 Germination depth vs. temperature requirements in naturally dispersed seeds of Leucospermum cordifolium and L. cuneiforme (proteaceae) G.J. Brits Vegetable and Ornamental Plant Research Institute, Private Bag X293, Pretoria, 0001 Republic of South Africa Submitted in partial fulfilment of an M.Sc. degree at the University of Stell en bosch (Botany Department) Accepted 9 October 1986 Myrmecochory (ant dispersal) and the requirement for both low and high diurnal soil temperatures during seed germination after fire, are important ecological determinants in the seedling recruitment phase of Leucospermum R. Br. species. A crucial unknown parameter linking these two factors is the effective burial depth (i.e. germination depth) of seeds stored in the subterranean seed bank. The germination depth of seedlings was effectively determined in a nursery trial by measuring the length of the hypocotyl between the radicle end and ground level. Using this method the germination depth of Leucospermum cordifolium (Salisb. ex Knight) Fourcade and L. cuneiforme (Burm. f.) Rourke seedlings emerging during winter, after summer burns in three localities was estimated. Average estimated germination depths varied between 30 and 45 mm. Daily minimum and maximum temperatures were measured at various soil depths during winter in burnt and unburnt mesic mountain fynbos. Temperatures measured at 10, 20 and 40 mm depths during winter were close to the known optimum daily low and high germination temperatures in L. cordifolium, in burnt fynbos, but not in unburnt fynbos. These temperatures in burnt fynbos prevailed especially during early winter. It was concluded that a high proportion of the Leucospermum cordifolium seed bank will germinate only during the first early winter following a fire. Other aspects of natural seed dispersal were studied. These included seedling dispersal numbers and patterns around burnt mother plants. Evidence for seed dispersal and burial by ants was found. Mierverspreiding van sade, asook die vereiste vir lae en hoe daaglikse grondtemperature gedurende kieming na 'n brand, is belangrike ekologiese determinante in die saad·regenerasie fase van Leucospermum R. Br. spesies. 'n Belangrike onbekende parameter wat hierdie faktore koppel, is die effektiewe diepte waarop sade begrawe word, d.i. die kiemingdiepte van sade wat in die ondergrondse saadbank geberg word. Die kiemingdiepte van saailinge is doeltreffend bepaal, in 'n kwekery-studie, deur die lengte van die hipokotiel tussen die kiemwortel­ kant en die grondoppervlak te meet. Met behulp van hierdie metode is die kiemingdiepte beraam van Leucospermum cordifolium (Salisb. ex Knight) Fourcade en L. cunei forme (Burm. f.) Rourke saailinge, wat in die winter na somer­ brande opgekom het in drie lokaliteite. Gemiddelde kiemingdieptes het gewissel van 30 tot 45 mm. Daaglikse minimum en maksimum temperature is op varierende gronddieptes gedurende die winter in gebrande en ongebrande mesiese berg-fynbos gemeet. Temperature wat op 10, 20 en 40-mm diepte gemeet is in die winter, was na aan die optimum daaglikse lae en hoe kiemingtemperature vir L. cordifolium, in gebrande fynbos, maar nie in ongebrande fynbos nie. Hierdie temperature in gebrande fynbos het veral gedurende die vroee winter geheers. Die afleiding is gemaak dat 'n hoe proporsie van die Leucospermum cordifolium-saadbank sal ontkiem slegs gedurende die eerste vroee winter wat volg na 'n brand. Ander aspekte van natuurlike saadverspreiding is bestudeer. Dit het die verspreidingsgetalle en -patrone van saailinge om verbrande moederplante ingesluit. Getuienis dat sade deur miere versprei en begrawe word, is gevind. Keywords: Fynbos, germination, Leucospermum, myrmecochory, seedling recruitment Introduction (i .e. germination depth) is, therefore, a crucial link between Seed regeneration in Leucospermum R. Br. is limited to the the hypotheses of myrmecochory and fluctuating temperature period immediately following fires, which occur naturally in requirement in germinating seeds. 6- to 4O-year cycles in the fynbos (Rourke 1972; Kruger This study examines seed dispersal and seedling recruitment 1979a). Daily fluctuating low and high temperatures are after fire in two Leucospermum species in relation to tem­ required for the germination of Leucospermum 'seeds' perature regimes at various soil depths in fynbos. The fol­ (achenes) and this requirement is met only in the immediate lowing problems are examined: post-fire environment during winter (Brits 1986a). (i) Can the depth at which seeds germinate be estimated? If Obligatory dependence of Leucospermum species on seed so, (ii) what is the relation between germination depth and dispersal and burial by ants in subterranean nests was pro­ the known temperature requirements of germinating seeds? posed by Bond & Slingsby (1983) and Slingsby & Bond (1985). (iii) Is there evidence that seeds were buried by ants and that In field experiments these authors recovered some buried seeds seedling recruitment depends on myrmecochory? from the upper galleries and channels of ant nests 4 to 7 cm below the soil surface. No quantitative study has been made Materials and Methods of the effective burial depth of Leucospermum seeds in the The relationship between hypocotyl length and depth subterranean seed bank, i.e. of seeds which contribute to the . of germination next generation of plants (viable seeds not buried too deep The emergence of germinating Leucospermum cordifolium or left under large stones). (Salisb. ex Knight) Fourcade seedlings was studied in a Daily temperature amplitude decreases sharply with in­ nursery. Freshly harvested L. cordifolium seeds were dis­ creasing germination depth of seeds (Mayer & Poljakoff­ infected, soaked in a 10,10 H202 solution and incubated at Mayber 1982). If seeds do not germinate at shallow depths, fluctuating temperatures (9°C for 16 h X 24°C for 8 h) in then the calculated temperature requirements for germination petri dishes (Brits 1986a). Germinating seeds were transplanted (Brits 1986a) may not be met. Effective seed burial depth as soon as the radicle had emerged into seedling trays 120 S.-Afr. Tydskr. Plantk. , 1987, 53(2) containing Riviersonderend mesic mountain fynbos soil (Brits burnt site: 125,11 103 lux 1986a). Before transplanting, the soil was flnned down slightly shaded site: 14,1 2 103 ± 7,53 103 lux. and allowed to 'settle' for approximately 3 weeks. Planting holes were made 10, 20 and 50 mm deep and seeds were Results and Discussion covered with wet soil. Ten seeds were planted at each depth. The relationship between hypocotyl length and depth Trays were placed in 50070 shade and irrigated once daily. of germination Seedling hypocotyls were marked at ground level after seed­ In species with epigeous seed germination, such as in Leu­ lings had emerged. On each seedling the distance was mea­ cospermum, the hypocotyl elongates in the soil, lifting the sured between the radicle end (foot) and the emergence level cotyledons from the point of germination to above the soil (ground level) of the hypocotyl. This distance was termed the surface (Foster & Gifford 1974). The hypocotyl in Leucosper­ hypocotyl length. mum is relatively stout, with a conspicuously flat, broadened, disc-like base (Van der Merwe 1977) or 'foot' from which Seed germination in fynbos the relatively thin radicle grows (Figure 1). In a field study the following three mesic mountain fynbos The mean length of hypocotyls at three planting depths communities (Moll et al. 1984), that had been burnt in were: accidental fires in the summer of 1984/85, were chosen for 10 mm planting depth: 10,6 ± 0,9 mm (9 out of 10 seedlings study during winter and spring 1985. emerged), (i) L. cordifolium, approximately eight-year-old (i.e. the time 20 mm planting depth: 21,8 ± 1,3 mm (4 out of 10 seedlings since the last fire), in the northern foothills of the Klein­ emerged), rivierberge near Tesselaarsdal (Caledon district, 34° 22'S, 19° 50 mm planting depth: 56,0 mm (lout of 10 seedlings 31'E). Some dominant proteoids of the area were Pro tea emerged). repens (L.) L., P. longifolia Andr. and Aulax umbellata (Thunb.) R.Br. The soil was a moderately deep sand over­ laying Table Mountain Group sandstone. The locality was burned on 27 December 1984. (ii) L. cordifolium, six-year-old, on a gentle south-facing slope, approximately 8 km from the sea near Elim (Bredas­ dorp district, 34° 40'S, 19° 37'E). The area, consisting of deep sand on Table Mountain Sandstone, was characterized by mixed stands of Protea compacta R. Br. and P. longifolia. It was burnt in January 1985. (iii) L. cuneiforme (Bunn. f.) Rourke, approximately ten-year­ old since fire, growing on sandy, shallow lithosolic soil on a south-facing, l3070 slope of the Riviersonderend mountain near Riviersonderend (Caledon district, 34° 9'S, 19° 54'E). The absolute age of these resprouter plants was not known. Other characteristic proteoid elements of the area were Leucadendron salignum Berg., Protea repens and dense stands of P. neriifolia R. Br. The locality was burned near the middle Figure 1 Hypocotyls of myrmecoehorous Leucospermum cordifolium of February 1985. (a); and serotinous Protea magnifica (b), P. compacta (c), and P. repens The burnt remains of Leucospermum plants as well as (d), 16 weeks after sowing at 10 mm depth. The base of each hypocotyl newly emergent seedlings were easily identifiable in each is indicated with an arrow. A = ground level. locality. The position of 5 or 6
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