The Spatial Signature of Biotic Interactions of a Clonal and a Non-clonal Palmetto in a Subtropical Plant Community

Spatial analyses of plant-distribution patterns can provide inferences about intra- and interspecific biotic interactions. Yet, such analyses are rare for clonal plants because effective tools (i.e., molecular markers) needed to map naturally occurring clonal individuals have only become available recently. Clonal plants are unique in that a single genotype has a potential to spatially place new individuals (i.e., ramets) in response to intra- and interspecific biotic interactions. Laboratory and greenhouse studies suggest that some clonal plants can avoid intra-genet, inter-genet, and inter-specific competition via rootplacement patterns. An intriguing and yet to be explored question is whether a spatial signature of such multi-level biotic interactions can be detected in natural plant communities. The facultatively clonal Serenoa repens and non-clonal Sabal etonia are ecologically similar and co-dominant palmettos that sympatrically occur in the Florida peninsula. We used amplified fragment length polymorphisms (AFLPs) to identify Serenoa genets and also to assign field-unidentifiable small individuals as Sabal seedlings, Serenoa seedlings, or Serenoa vegetative sprouts. Then, we conducted univariate and bivariate multi-distance spatial analyses to examine the spatial interactions of Serenoa (n=271) and Sabal (n=137) within a 20x20 m grid at three levels, intragenet, intergenet and interspecific. We found that spatial interactions were not random at all three levels of biotic interactions. Serenoa genets appear to spatially avoid self-competition as well as intergenet competition. Furthermore, Serenoa and Sabal were spatially negatively associated with each other. However, this negative association pattern was also evident in a spatial comparison between non-clonal Serenoa and Sabal, suggesting that Serenoa genets’ spatial avoidance of Sabal through placement of new ramets is not the explanation of the interspecific-level negative spatial pattern. Our results emphasize the importance of investigating spatial signatures of biotic as well as abiotic interactions at multiple levels in understanding spatial distribution patterns of clonal plants in natural plant communities.

Provenance signature of a forearc basin modified by spreading ridge subduction: Detrital zircon geochronology and detrital modes from the Paleogene Arkose Ridge Formation, southern Alaska

Upper Paleocene–Eocene boulder conglomerate, cross-stratified sandstone, and laminated carbonaceous mudstone of the Arkose Ridge Formation exposed in the southern Talkeetna Mountains record fluvial-lacustrine deposition proximal to the volcanic arc in a forearc basin modified by Paleogene spreading ridge subduction beneath southern Alaska. U-Pb ages of detrital zircon grains and modal analyses were obtained from stratigraphic sections spanning the 2,000 m thick Arkose Ridge Formation in order to constrain the lithology, age, and location of sediment sources that provided detritus. Detrital modes from 24 conglomerate beds and 54 sandstone thin sections aredominated by plutonic and volcanic clasts and plagioclase feldspar with minor quartz, schist, hornblende, argillite, and metabasalt. Westernmost sandstone and conglomerate strata contain <5% volcanic clasts whereas easternmost sandstone and conglomerate strata contain 40 to >80% volcanic clasts. Temporally, eastern sandstones andconglomerates exhibit an upsection increase in volcanic detritus from <40 to >80% volcanic clasts. U-Pb ages from >1400 detrital zircons in 15 sandstone samples reveal three main populations: late Paleocene–Eocene (60-48 Ma; 16% of all grains), Late Cretaceous–early Paleocene (85–60 Ma; 62%) and Jurassic–Early Cretaceous (200–100 Ma; 12%). A plot of U/Th vs U-Pb ages shows that >97% of zircons are <200 Ma and>99% of zircons have <10 U/Th ratios, consistent with mainly igneous source terranes. Strata show increased enrichment in late Paleocene–Eocene detrital zircons from <2% in the west to >25% in the east. In eastern sections, this younger age population increases temporally from 0% in the lower 50 m of the section to >40% in samples collected >740 m above the base. Integration of the compositional and detrital geochronologic data suggests: (1) Detritus was eroded mainly from igneous sources exposed directly north of the Arkose Ridge Formation strata, mainly Jurassic–Paleocene plutons and Paleocene–Eocenevolcanic centers. Subordinate metamorphic detritus was eroded from western Mesozoic low-grade metamorphic sources. Subordinate sedimentary detritus was eroded from eastern Mesozoic sedimentary sources. (2) Eastern deposystems received higher proportions of juvenile volcanic detritus through time, consistent with construction of adjacent slab-window volcanic centers during Arkose Ridge Formation deposition. (3)Western deposystems transported detritus from Jurassic–Paleocene arc plutons that flank the northwestern basin margin. (4) Metasedimentary strata of the Chugach accretionaryprism, exposed 20-50 km south of the Arkose Ridge Formation, did not contribute abundant detritus. Conventional provenance models predict reduced input of volcanic detritus to forearc basins during exhumation of the volcanic edifice and increasing exposure ofsubvolcanic plutons (Dickinson, 1995; Ingersoll and Eastmond, 2007). In the forearc strata of these conventional models, sandstone modal analyses record progressive increases upsection in quartz and feldspar concomitant with decreases in lithic grains, mainly volcanic lithics. Additionally, as the arc massif denudes through time, theyoungest detrital U-Pb zircon age populations become significantly older than the age of forearc deposition as the arc migrates inboard or ceases magmatism. Westernmost strata of the Arkose Ridge Formation are consistent with this conventional model. However, easternmost strata of the Arkose Ridge Formation contain sandstone modes that record an upsection increase in lithic grains accompanied by a decrease in quartz and feldspar, and detrital zircon age populations that closely match the age of deposition. This deviation from the conventional model is due to the proximity of the easternmost strata to adjacent juvenile volcanic rocks emplaced by slab-window volcanic processes. Provenance data from the Arkose Ridge Formation show that forearc basins modified by spreading ridge subduction may record upsection increases in non-arc, syndepositional volcanic detritusdue to contemporaneous accumulation of thick volcanic sequences at slab-window volcanic centers. This change may occur locally at the same time that other regions of the forearc continue to receive increasing amounts of plutonic detritus as the remnant arc denudes, resulting in complex lateral variations in forearc basin petrofacies and chronofacies.