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Plant Wax Alkanes and Alcohols As Herbivore Diet Composition Markers

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Plant Wax Alkanes and Alcohols As Herbivore Diet Composition Markers J. Range Manage. 57:259 -268 May 2004 Plant wax alkanes and alcohols as herbivore diet composition markers MIGUEL N. BUGALHO, HUGH DOVE, WALTER KELMAN, JEFF T. WOOD, AND ROBERT W. MAYES Bugalho is a post- doctoral Research Fellow at Centro de Ecologia Aplicada, Instituto Superior de Agronomia, 1349 -017 Tapada da Ajuda, Lisboa, Portugal; Dove and Kelman are Senior Researchers at CSIRO Plant Industry, GPO Box 1600, Canberra, ACT 2601, Australia; Wood is a statistician who was at CSIRO Mathematical and Information Sciences, GPO Box 664, Canberra, ACT 2601, Australia during the work and presently is at the Statistical Consulting Unit, John Dedman Building, Australian National University, ACT 0200, Australia; Mayes is a Senior Researcher at The Macaulay Institute, Craigiebuckler, Aberdeen AB1 S 8QH, Scotland. Abstract Resumen The n- alkanes in plant cuticular wax have been used as mark- Los n- alcanos de la cera cuticular de las plantas han sido usa- ers for estimating the species composition of herbivore diets, but dos como marcadores para determinar la composición de the long -chain fatty alcohols (LCOH) of plant wax may also be especies en las dietas de los herbívoros, pero los alcoholes grasos useful. The objective of this research was to assess if LCOH con- de cadena larga (LCOH) también pueden ser útiles. El objetivo tributed extra information to differentiate plant species, com- de esta investigación fue evaluar si los LCOH aportan informa- pared with n- alkanes only. We used 3 data sets consisting of n- ción extra para diferenciar las especies de plantas, comparados alkane and LCOH concentrations of plant species occurring in con los n- alcanos solos. Usamos 3 juegos de datos consistentes de pastures of New South Wales, Australia. We used Principal concentraciones de n- alcanos y LCOH de especies de plantas que Component Analysis (PCA) to summarise the data for n- alkane están presentes en los potreros de Nueva Gales del Sur, and LCOH concentrations obtained for the species in these data Australia. Usamos el análisis de componentes principales (PCA) sets. The first 3 principal components explained 86 to 93% and para resumir los datos de las concentraciones de n- alcanos y 75 to 99% of the variance in n- alkane and LCOH concentrations, LCOH obtenidos para las especies de plantas de los juegos de respectively. Orthogonal Procrustes Rotation (OPR) was then datos utilizados. Los tres primeros componentes principales used to compare the results of PCA conducted with n- alkane and explicaron del 86 al 93% y del 75 al 99% de la variación en las LCOH data, with a view to establishing whether LCOH provided concentraciones de n- alcanos y LCOH respectivamente. Entonces discriminatory information in addition to that provided by the n- se uso la Rotación de Procrustes Ortogonales (OPR) para com- alkanes. Results of OPR indicated that this was so for all 3 data parar los resultados de los componentes principales realizados con sets, and suggested that the LCOH would be useful additional los datos de n- alcanos y LCOH, con la mira de establecer si los markers for discriminating between plant species. We tested this LCOH proveyeron información discriminatoria adicional a la by using Discriminant Analysis and cross - validation procedures proporcionada por los n- alcanos. Los resultados de OPR indi- in 2 data sets to distinguish between defined species groups of C3 caron que esto si fue así para los 3 juegos de datos y sugirieron que grasses, C4 grasses, clovers and Lotus spp. The discrimination los LCOH pudieran ser marcadores adicionales útiles para dis- between these categories and the proportion of plant species cor- criminar entre especies de plantas. Probamos esto usando proced- rectly classified into the defined categories was better when using imientos de Análisis Discriminante y validación cruzada en dos n- alkanes and LCOH together, compared with alkanes alone. juegos de datos para distinguir entre grupos de especies definidas Our results indicate that LCOH provided additional information de zacates C3, zacates, C4, tréboles y Lotus spp. La discriminación that could be used for distinguishing plant species as part of esti- entre estas categorías y la proporción de especies de plantas cor- mating the species composition of herbivore diets. rectamente clasificadas dentro de las categorías definidas fue mejor cuando se uso n- alcanos y LCOH juntos en comparación Key Words: n- alkanes, long -chain fatty alcohols, markers, diet con los alcanos solos. Nuestros resultados indican que LCOH proveyeron información adicional que pudiera ser utilizada para composition, herbivores distinguir especies de plantas como parte de la estimación de la composición de especies de las dietas de los herbívoros. The species composition of herbivore diets can be estimated by exploiting differences among plant species in their cuticular wax compounds, especially n- alkanes (Dove and Mayes 1991). The n- alkanes are relatively inert, long -chain saturated hydrocarbons(C21 to C37) that, in ruminants, can be recovered in feces to a degree dependent on their carbon -chain length (Dove and Mayes 1991, 1996). Diet composition is estimated by relating the n -alka- Miguel Bugalho thanks the Portuguese Ministry of Science and Technology (Post- ne content of feces to the n- alkane content of the plants available doctoral grant FCT /SFRH/727 /2000) and CSIRO Plant Industry, Australia, for to herbivores, after correcting for n- alkane fecal recovery (e.g. support during a period as Visiting Scientist at CSIRO. We thank Denys Garden Dove and Moore 1995). Using plant wax n- alkanes or other wax and David Officer of NSW Agriculture, Australia, for the provision of the herbage samples in data sets 2 and 3, respectively. We also thank Stuart Lamb and Susan components as diet markers offers several advantages over other Hoebee for their assistance with the n- alkane and LCOH analyses and Christine nutritional markers (Dove and Mayes 1999, Mayes and Dove Donnelly for the preparation of Figures 1 to 3. 2000, Sandberg et al. 2000). An important constraint, however, is Manuscript accepted 17 Aug. 03. JOURNAL OF RANGE MANAGEMENT57(3) May 2004 259 that the number of species that can be dif-57' E (Data set 3). Differences in theN- alkane and LCOH extraction and ferentiated in the diet cannot exceed the method of sample collection and the nature quantification number of alkane markers available; thisof the material collected suggest that these The n- alkanes in data sets 1 and 2 were limits differentiation to usually lower than may be regarded as 3 independent dataextracted and quantified as described by 15 species (Mayes and Dove 2000). When sets, although combinations of data setsMayes et al. (1986), with modifications dealing with complex plant communitieswere also examined to explore the trendsindicated by Salt et al. (1992); internal such as rangelands, the number of plant in between - species differences in the pat- standards used were docosane (C22 alka- species available to the herbivore may terns of alkane or LCOH concentrations. ne) and tetratriacontane (C34 alkane). In thus exceed the number of n- alkane mark- Data set 1 consisted of shoot materialaddition, 1- pentacosanol (C25 normal ers available. One means of overcomingfrom 5 grass species (Austrodanthoniaalcohol) was used as an internal standard this constraint is to include more classes ofrichardsonii (Cashmore) H.P. Linder cv. to quantify the LCOH. It should be noted plant wax components as markers for dis- Taranna, Phalaris aquatica L., Vulpia that the alkaline extraction conditions used criminating between plant species. A pre- myuros (L.) Gmel., Bromus catharticusfor the n- alkanes (Mayes et al. 1986) requisite of this approach is that the addi-Vahl. and Festuca arundinacea Schreb.) extract not only the free LCOH of cuticu- tional markers must provide discriminato- and 7 legumes (Trifolium striatum L., T. lar wax, but also the LCOH bound as wax ry information additional to that providedglomeratum L., T. repens L., Lotus comic -esters. Extracted LCOH were eluted from by the n- alkanes. ulatus L. "Prostrate ", L. corniculatus L. silica gel columns using 1:1 heptane:ethanol Long -chain fatty alcohols (LCOH) arecv. Goldie, L. pedunculatus Cay. cv. immediately following alkane elution. The widespread in the plant species consumed Sharnae, L. pedunculatus Cay. cv.LCOH eluates were dried, redissolved in by herbivores and frequently occur atGrasslands Maku). Individual plants of 1:1 heptane:ethanol and injected into a 30m higher concentrations than n- alkaneseach species were cut from a grazed sward x 0.53mm ID BPX35 bonded -phase silica (Kolattukudy 1976, Dove and Mayes(Kelman et al. 2003) using electric clip-column (0.5mm film thickness; SGE Pty 1999, Mayes and Dove 2000, Kelman etpers, and were then composited within Ltd Australia) in a PU4550 gas chromato- aí.2003). Furthermore, the LCOH can be species to provide sufficient material for graph fitted with a splitless injector and separated and quantified by an extensionplant wax analyses. flame ionization detector. The carrier gas (unpublished data, R.W. Mayes) of the The second data set consisted of leaf tis- was helium. existing procedure used to analyse n -alka- sue from 7 grass species (Austrodanthonia Because of difficulties encountered in nes. This suggests that LCOH could beracemosa (R. Br.) H.P. Linder, A. pilosa separating alcohols from sterols under the useful diet composition markers in addi- (R. Br.) H.P. Linder, Elymus scaber (R. chromatography conditions used, it is possi- tion to the n- alkanes. Br.) A. Love, Bothriochloa macra (Steud.) ble that the compounds described as 1 -C27- Our objectives in the present studyS.T. Blake, Chloris gayana Kunth, OL and 1-C29-0L could in fact be sterols. were: Themeda australis (R. Br.) Stapf., However, as this does not affect subsequent 1. To quantify n- alkane and LCOH con-Microlaena stipoides (Labill.) R. Br.) with data analysis or the arguments presented in centrations in the cuticular wax of pas-the last species coming from 2 samplingthis paper, these components are assumed ture species commonly found in Newlocationsatthesite(designated hereafter to be primary alcohols.
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