Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures

Aim In a selected literature survey we reviewed studies on the habitat heterogeneity–animal species diversity relationship and evaluated whether there are uncertainties and biases in its empirical support. Location World-wide. Methods We reviewed 85 publications for the period 1960–2003. We screened each publication for terms that were used to define habitat heterogeneity, the animal species group and ecosystem studied, the definition of the structural variable, the measurement of vegetation structure and the temporal and spatial scale of the study. Main conclusions The majority of studies found a positive correlation between habitat heterogeneity/diversity and animal species diversity. However, empirical support for this relationship is drastically biased towards studies of vertebrates and habitats under anthropogenic influence. In this paper, we show that ecological effects of habitat heterogeneity may vary considerably between species groups depending on whether structural attributes are perceived as heterogeneity or fragmentation. Possible effects may also vary relative to the structural variable measured. Based upon this, we introduce a classification framework that may be used for across-studies comparisons. Moreover, the effect of habitat heterogeneity for one species group may differ in relation to the spatial scale. In several studies, however, different species groups are closely linked to ‘keystone structures’ that determine animal species diversity by their presence. Detecting crucial keystone structures of the vegetation has profound implications for nature conservation and biodiversity management.     

Plant light interception can be explained via computed tomography scanning: demonstration with pyramidal cedar (Thuja occidentalis, Fastigiata)

BACKGROUND AND AIMS: Light interception by the leaf canopy is a key aspect of plant photosynthesis, which helps mitigate the greenhouse effect via atmospheric CO(2) recycling. The relationship between plant light interception and leaf area was traditionally modelled with the Beer-Lambert law, until the spatial distribution of leaves was incorporated through the fractal dimension of leafless plant structure photographed from the side allowing maximum appearance of branches and petioles. However, photographs of leafless plants are two-dimensional projections of three-dimensional structures, and sampled plants were cut at the stem base before leaf blades were detached manually, so canopy development could not be followed for individual plants. Therefore, a new measurement and modelling approach were developed to explain plant light interception more completely and precisely, based on appropriate processing of computed tomography (CT) scanning data collected for developing canopies. METHODS: Three-dimensional images of canopies were constructed from CT scanning data. Leaf volumes (LV) were evaluated from complete canopy images, and fractal dimensions (FD) were estimated from skeletonized leafless images. The experimental plant species is pyramidal cedar (Thuja occidentalis, Fastigiata). KEY RESULTS: The three-dimensional version of the Beer-Lambert law based on FD alone provided a much better explanation of plant light interception (R(2) = 0.858) than those using the product LV*FD (0.589) or LV alone (0.548). While values of all three regressors were found to increase over time, FD in the Beer-Lambert law followed the increase in light interception the most closely. The delayed increase of LV reflected the appearance of new leaves only after branches had lengthened and ramified. CONCLUSIONS: The very strong correlation obtained with FD demonstrates that CT scanning data contain fundamental information about the canopy architecture geometry. The model can be used to identify crops and plantation trees with improved light interception and productivity.     

Estimation of plant and leaf area index using three techniques in a mature native eucalypt canopy

Abstract Leaf area index (L) is a critical variable in monitoring and modelling forest condition and growth and is therefore important for foresters and environmental scientists to measure routinely and accurately. We compared three different methods for estimating L: a plant canopy analyser (PCA), a point‐quadrat camera method and digital hemispherical photography at a native eucalypt forest canopy at Tumbarumba in southern New South Wales, Australia. All of these methods produced indirect estimates of L based on the close coupling between radiation penetration and canopy structure. The individual L estimates were compared, and the potential advantages and disadvantages of each method were discussed in relation to use in forest inventory and in field data collection programmes for remote sensing calibration and verification. The comparison indicated that all three methods, PCA, digital hemispherical photography and the modified point‐quadrat camera method, produced similar estimates with a standard error between techniques of less than 0.2 L units. All methods, however, provided biased estimates of L and calibration is required to derive true stand L. A key benefit, however, of all of these estimation methods is that observations can be collected in a short period of time (1–2 h of field‐work per plot).     

Direct somatic embryogenesis and plant regeneration from leaf, petiole, and stem explants of Golden Pothos

Somatic embryos directly formed at cut edges or on the surface of leaf explants, around cut ends or along side surfaces of petiole and stem explants of Golden Pothos [Epipremnum aureum (Linden & Andre) Bunt.] on Murashige and Skoog (MS) medium supplemented with N-(2-chloro-4-pyridyl)-N-phenylurea (CPPU) or N-phenyl-N-1, 2, 3-thiadiazol-5-ylurea (TDZ) with -naphthalene acetic acid (NAA) and a medium called MK containing MS salts with Kaos vitamins, supplemented with 2.0 mg/l TDZ and 0.2 mg/l NAA. Somatic embryos were also produced on MS medium containing 2.0 mg/l kinetin (KN) and 0.5 mg/l 2,4-dichlorophenoxyacetic acid (2,4-D) from leaf and petiole explants, MS medium supplemented with 2.0 mg/l CPPU and 0.5 mg/l 2,4-D from petiole and stem explants, and 2.0 mg/l TDZ and 0.2 mg/l or 0.5 mg/l 2,4-D from stem explants. In addition, somatic embryos occurred from stem explants on Chus N₆ medium containing 2.0 mg/l CPPU and 0.2 mg/l NAA. Somatic embryos matured and grew into multiple buds, shoots, or even plantlets after 2–3 months on the initial culture medium. Germination was optimal on MS medium containing either 2 mg/l 6-benzylaminopurine (BA) and 0.2 mg/l NAA or 2 mg/l zeatin and 0.2 mg/l NAA. Shoots elongated better and roots developed well on MS medium with no growth regulators. Approximately 30–100 plantlets were regenerated from each explant. The regenerated plants grew vigorously after transplanting to a soil-less container substrate in a shaded greenhouse.     

Vegetative and reproductive structure of the extinct Platanus neptuni from the Tertiary of Europe and relationships within the Platanaceae

Platanus neptuni (Ettingshausen) Bek, Holý & Z. Kvaek was a conspicuous warm-temperate to subtropical element of Late Eocene to Late Miocene European floras. In our concept, the P. neptuni plant includes not only globose infructescences upon which the species epithet is based, but also staminate and pistillate inflorescences and distinctive stipulate foliage. The leaves range from simple (P. neptuni morphoforma reussii (Ett.) comb. et stat. nov.) to trifoliolate (P. neptuni morphoforma fraxinifolia (Johnson & Gilmore) comb. et stat. nov.) and sometimes quinquefoliolate (P. neptuni morphoforma hibernica (Johnson & Gilmore) comb. et stat. nov.) with unlobed elliptical to obovate laminae that are uniform in venation, marginal serration, and epidermal structure. Foliar twigs confirm that the leaves are deciduous, with each petiole base enveloping a bud, as in extant Platanus subgen. Platanus. Platanus neptuni differs from extant species of the genus by large peltate glandular trichomes on the fruits and leaves, a prominent circumscissile rim on the stalk below the inflorescence, as well as by the tendency for compound foliage. These characters justify its position within an extinct subgenus of the Platanaceae (Platanus L. subgen. Glandulosa Z. Kvaek, Manchester & Guo). Platanus neptuni was common in mesic humid subtropical forests on volcanogenic subtrates and at sea shores.     

Spatial distribution of photosynthetic response to long-term influence of elevated CO2 in a mediterranean macchia mini-ecosystem

During an open-top chamber experiment performed in evergreen ‘macchia’ ecosystem, which was represented by the clumps of natural vegetation dominated by Quercus ilex trees, the trees were exposed to one of two CO₂ concentrations (ambient CO₂, AC-variant and elevated CO₂, i.e. ambient plus 350 μmol CO₂ mol⁻¹, EC-variant) continuously over 5 years. Clumps of natural vegetation were enclosed in open top chambers (OTCs). Within the crowns of investigated Quercus ilex trees in OTCs, two crown layers i.e. sunny (E-leaves) and shaded (S-leaves), were identified as differing in solar radiation environment. To evaluate the effect of elevated CO₂, as well as the functional differentiation in assimilation activity of E- versus S-leaves, gas exchange and chlorophyll fluorescence techniques were used. The stimulatory effect of the long-term elevated CO₂ on the A_N–PPFD relation was evident in E- and S-leaves of investigated Quercus trees. The A_Nmax sensitivity of AC-variant leaves to the sudden application of elevated CO₂ was higher for S-leaves (42%) than for E-leaves (24%). The PPFD saturated rate of regulated thermal energy dissipation (ERD) confirmed the foliage differentiation caused by the long-term influence of elevated CO₂. The ERD of E- and S-leaves in the AC-variant were 1.11 times of that in the EC-variant. However, the estimated rates of photochemistry (ERP) of E- and S-leaves in the EC-variant were 1.35 and 1.22 times of E- and S-leaves in the AC-variant. The achievement of the critical value of q_P=0.4 in E- and S-leaves from the EC-variant under lower values of PPFD compared to the AC-variant indicates a higher degree of PSII over-reduction. Thus, elevated CO₂ can be responsible for an increased susceptibility of photosynthetic apparatus to high irradiance. The obtained results support the hypothesis on the foliage vertical distribution effect on the whole canopy response to elevated CO₂. The S-layers of the canopy can play an important role in providing storage space for photosynthesis under elevated CO₂.     

Variation in the δ13C of foliage of Pinus sylvestris L. in relation to climate and additions of nitrogen: analysis of a 32-year chronology

We report an analysis of both the long‐ and short‐term drivers of the carbon (C) isotope composition (δ¹³C) values of current year needles of Pinus sylvestris L. linked to changing atmospheric carbon dioxide (CO₂) concentrations (c_a) and climate using data from a uniquely long‐term nitrogen (N) fertilization experiment in the north of Sweden (consisting of three N dosage levels and a control treatment) from 1970 until 2002. N loading produced trees with less negative δ¹³C of foliage, by around 0.45‰ on average, with the difference in δ¹³C between control and N treatments not dependant upon N dosage. The average δ¹³C values decreased at a rate of around 0.03‰ yr⁻¹, even after accounting for the Suess effect (the decrease in the atmospheric CO₂δ¹³C due to anthropogenic emissions of isotopically light CO₂). This decrease is large enough to cause a significant, progressive change in the δ¹³C down through a soil profile. Modelled values of plant intrinsic water use efficiency (WUE_i) and the ratio of leaf internal to external [CO₂] (c_i/c_a) showed that this was the result of c_i increasing in parallel with c_a (while c_i/c_a increased), thus causing little change in WUE_i over the 32 years of study. The residuals from the relationships between year and δ¹³C were used to examine the impact of climate on the interannual variation of C isotope composition of needles. This included the use of a fire hazard index (FHI) model, which integrates climatic factors known to influence plant stomatal conductance and hence δ¹³C. The FHI produced the best fit with δ¹³C values when climate data were averaged over the whole growth season (for control plots) and for July for all the N treatments, explaining ca. 60% of the total interannual variation in δ¹³C. Further, trees from the N treatments appeared more susceptible to air‐humidity‐based climate parameters, as seen from higher correlation coefficients, than were control trees. Thus, our data suggest the possibility of increased susceptibility to drought conditions in ecosystems with moderate to high N deposition rates. Also, there is the possibility that, because there was no apparent change in WUE_i of P. sylvestris in this ecosystem over the last 32 years, the rate of sequestration of C into boreal ecosystems may not increase with c_a, as has been predicted.     

Feed resource selection of Criollo goats artificially infected with Haemonchus contortus: nutritional wisdom and prophylactic self-medication

Previous cafeteria studies suggested that a moderate natural gastrointestinal nematode (GIN) infection did not modify the resource selection of adult Criollo goats towards tannin-rich plants compared with worm-free goats. A higher infection with Haemonchus contortus could trigger a change in the resource selection behaviour towards tannin-rich foliage. Alternatively, goats might select plant species solely to meet their nutritional requirements. A cafeteria study investigated the effect of a high artificial infection with H. contortus on the feed resource selection of goats. Adult Criollo goats (37.5±4.8 kg BW) with browsing experience were distributed in two groups: the infected group (IG) with six animals artificially infected with H. contortus (6000 L₃/animal); and the non-infected group (NIG) with six animals maintained worm-free. The experiment included two 5-day periods with additional 5-day adaptation period. In the first period, animals were offered foliage of five plant species with a decreasing gradient of condensed tannins (CT) (Mimosa bahamensis, Gymnopodium floribundum, Havardia albicans, Acacia pennatula, Lysiloma latisiliqum), and three plant species with negligible CT content (Leucaena leucocephala, Piscidia piscipula and Brosimum alicastrum). In the second period the foliage of B. alicastrum was withdrawn. A grain-based concentrate feed was offered daily at 1% BW in DM basis. Dry matter and nutrient intake was determined. Foliage selection of each experimental group was determined using the Chesson selection index. The H. contortus egg count per gram of faeces (EPG) was determined for infected goats twice daily. Chesson index showed a similar pattern of foliage selection on periods 1 and 2. Mean EPG of goats in IG was 2028±259 EPG during period 1 and 1 293±198 EPG during period 2 (P>0.05). During period 1, the selection pattern was highest for B. alicastrum (tannin-free), followed by a tannin-rich plant (M. bahamensis). These two plants remained as highly selected during period 2. The Chesson index showed that both experimental groups (IG and NIG) selected the same plant species in both periods. Thus, a high H. contortus infection did not affect selection of goats fed with CT-rich plants. Apparently, goats balanced their nutrient intake with the plants selected, showing evidence of nutritional wisdom. This balance may have helped to prevent excess protein in the diet and also to maintain a low GIN infection, both considered as examples of prophylactic self-medication.     

Inferring changes in foliar mass and area from foliage cover: A mechanistic model

Measuring foliar area or mass directly is destructive, and precludes long‐term, repeated observations of individual trees as they suffer or recover from foliar damage. Instead, foliage cover indices are often used as a proxy for foliar mass. Patterns of fluctuations in foliage cover indices can be used to infer qualitative changes in canopy health. However, foliage cover is not necessarily linearly related to foliar area or mass, and this may confound the detection of significant foliar damage, and comparisons of herbivore browse impacts between individual trees, tree species or sites. I derived a mechanistic model to quantify the relationship between foliar area or mass and foliage cover measured as the proportion of sky occluded by leaves. This one parameter model is close to linear for single‐tiered trees, but increasingly non‐linear for multi‐tiered trees. I compared the non‐linear model to a linear model using foliage cover data from an artificial defoliation experiment on two single‐tiered, sub‐canopy species and from simulated photographic images of single‐ and multi‐tiered canopies. The non‐linear model had lower errors than the linear model, and errors did not increase with foliage density (leaf area per unit area), variation (of leaf sizes within and between canopies) or leaf geometry. The non‐linear model can be easily parameterized from relatively low‐cost observations of foliage cover, independently of empirical measurements of foliar area or mass, and is applicable to a wide range of tree species. It should therefore help managers quantify how changes in foliage cover due to natural fluctuations or foliar damage affect foliar area and mass, and can be used to quantify parameters for models of browse impacts in mixed forest.