Genetic diversity,differentiation and conservation in <Emphasis Type="Italic">Araucaria bidwillii</Emphasis>(Araucariaceae), Australia's Bunya pine

Habitat fragmentation has resulted in many species becoming geographically restricted, as dispersal among subsequent isolates becomes compromised. This study investigated the effects of historical fragmentation on the genetic diversity and differentiation of Australia's Bunya Pine (Araucaria bidwillii) using random amplified DNA (RAPD) markers. High diversity characterises all Bunya populations sampled, regardless of population size or degree of isolation, which may be either due to similar effective population size among populations, or the lagging effects of the detection of contemporary genetic signal in long-lived conifer species. Large genetic differentiation characterises the northern and southern populations, although all sampled populations are significantly differentiated from each other. The northern population, at Mt Lewis, is responsible for the majority of variation detected among populations, and as such represents a significant genetic reservoir. The conservation of the genotypes of the Mt Lewis population may be imperative for the future of the species, given predicted models of accelerated climate change over the coming decades.     

High-throughput DNA extraction from forest trees

It is difficult to extract pure high-quality DNA from trees, which may not be amenable to advances in extraction methods suitable for other plants. A new commercial high-throughput DNA extraction system, using a silica binding matrix for purification and a multisample mixer mill for tissue disruption, was evaluated for its suitability withEucalyptus spp.,Pinus spp., andAraucaria cunninghamii (hoop pine). DNA suitable for a range of molecular biology applications was successfully extracted from all genera. The method was highly reliable when tested in more than 500 preparations and could be adapted to different tree species with relatively minor modifications.     

Anomalous ring identification in two Australian subtropical Araucariaceae species permits annual ring dating and growth-climate relationship development

Almost all Australian tropical and subtropical regions lack annually-resolved long-term (multi-decadal to centennial scale) instrumental climate records. Reconstructing climate in these regions requires the use of sparse climate proxy records such as tree rings. Tree rings often archive annually-resolved centennial-scale climate information. However, many tropical and subtropical species have short life-spans, the timbers are poorly preserved, and there is a belief that the proxy records of these species are often compromised by ring anomalies. Additionally, for many species the relationship between climate (e.g. temperature and/or rainfall) and tree growth has not been established. These factors have led to tree-ring data being underutilized in the Australian subtropics. Trees in the Araucariaceae family, a common family in northern and eastern Australia, are both longer lived than many species in the Australian subtropics, present growth rings that are annual in nature, and their growth is known to vary with climate. In this study we examine two subtropical Araucariaceae species, Araucaria cunninghamii and Araucaria bidwillii, and quantify the relationship between their radial growth and climate variability. Ring anomalies including false, faint, locally absent, and pinching rings, are found to be present in these species, however, bomb-pulse radiocarbon dating of A. cunninghamii samples together with a whole tree approach helped to identify annual growth patterns despite such anomalous ring boundaries. Additionally, to determine which climate variables most influence growth in these species, dendrometers were installed at two locations in subtropical Southeast Queensland, Australia. We found that rainfall variability drives annual ring growth, while temperature constrains the onset and conclusion of the growth season each year. Our results demonstrate that through the use of A. cunninghamii and A. bidwillii trees which demonstrate annual growth in relation to climate variables there is potential to develop centennial scale climate reconstructions from the Australian subtropics. We provide recommendations on how to best identify ring anomalies in these species to help in the future development of long-term chronologies and climate reconstructions.     

Salinity and Temperature Effects on Germination for Three Salt-resistant Euhalophytes, Halostachys caspica, Kalidium foliatum and Halocnemum strobilaceum

The effects of three salinities (0, 100 and 500 mM NaCl) and four constant temperatures (10, 20, 30 and 35 °C) on seed germination of Halostachys caspica (M. B.) C. A. Mey., Kalidium foliatum (Pall.) Mop. and Halocnemum strobilaceum (Pall.) Bieb. were investigated. After seeds were treated with different concentrations of NaCl at constant temperatures of 10–35 °C for 16 days, ungerminated seeds were transferred to distilled water for 10 days to investigate the total germination; after this time, the ungerminated seeds from the 10 and 20 °C treatments were then moved to 35 °C for another 5 days to determine the final germination. The three plant species in the present experiment are salt-resistant euhalophytes growing in high saline soils in the Zhungur Basin in Xinjiang, a northwest province of China.Compared with germination under control conditions, germination percentages of all three species were not affected by 100 mM NaCl at 10–35 °C, while severely inhibited by 500 mM NaCl; germination percentages were very low at 10 °C up to 100 mM NaCl for all species; the optimum temperature for germination of H. caspica and K. foliatum was 20–30 °C, while 35 °C for H. strobilaceum, up to 100 mM NaCl; seeds did not suffer ion toxicity for all species, as evidenced by the high total germination after ungerminated seeds pretreated with 500 mM NaCl were transferred to distilled water at constant temperatures of 10–35 °C for 10 days, and the high final germination after the ungerminated seeds from the 10 and 20 °C treatments were subsequently moved to 35 °C for another 5 days; Halostachys caspica had greater sensitivity to increasing temperatures from 10 and 20 °C to 35 °C compared with the other two species.     

Germination-associated events and the desiccation sensitivity of recalcitrant seeds — a study on three unrelated species

The storage behaviour of recalcitrant seeds was assessed using three diverse species: a gymnosperm, Araucaria angustifolia (Bert.) O. Kuntze; a herbaceous monocotyledon, Scadoxus membranaceus (Bak.) Friis Nordal; and a woody dicotyledon, Landolphia kirkii Dyer. Seeds were stored under conditions of high relative humidities that maintained seed moisture content and under low relative humidities that caused drying. At regular intervals moisture content was determined, germinability assessed and the ultrastructure of radicle meristem cells examined. Under storage at high relative humidity, seed moisture content was maintained at the original level and subcellular germination events were initiated in the short-term. Such seeds showed enhanced rates of germination when removed from storage and planted. Long-term storage under these conditions resulted in the initiation of subcellular damage which intensified with time and ultimately resulted in the loss of viability. The rate at which germination events proceeded varied among the three species, and could be directly correlated with the period of viability retention under humid storage conditions. Storage under desiccating conditions resulted in subcellular damage and rapid loss of viability. The rate at which the seeds dried varied among the three species. The proportion of water loss tolerated by the different species before loss of viability, correlated with the rate of drying. The storage behaviour of the seeds of these three species is discussed in terms of a previously described model.     

The Effect of Desiccation on the Longevity of Seeds of Araucaria hunsteinii and A. cunninghamii

Seeds of Araucaria hunsteinii K. Schum. dried quicker at 29°Cthan at 19°C and quicker with the seed-coat removed thanwhen intact; seeds enclosed in polyethylene bags increased inmoisture content. At 15°C, seeds in a flow of air driedquicker than seeds in a box with silica gel, which in turn driedquicker than seeds in a box with no desiccant. No loss of germinationability occurred on drying fresh seed from 53 to about 32 percent moisture content (fresh weight basis); during further desiccationthe percentage germination was related to percentage moisturecontent in the form of a sigmoid curve, culminating in a completefailure to germinate at approximately 14 per cent moisture content.A consistent relationship was observed for all treatments andthe mean critical moisture content for seed death (failure togerminate) was near 23 per cent. Excised embryos grew on 1 percent agar but died if previously subjected to 14 h of desiccationat 15°C. In contrast, no relationship was found between germination andmoisture content of A. cunninghamii D. Don on desiccation from21 to 7 per cent moisture content. Possible causes for the observeddifference in response to desiccation are discussed and methodsfor seed storage are considered. Araucaria hunsteinii, Klinkii pine, Araucaria cunninghamii, Hoop pine, desiccation, seed longevity, storage of seeds     

Woody species diversity influences productivity and soil nutrient availability in tropical plantations

We investigated the relationship between plant diversity and ecological function (production and nutrient cycling) in tropical tree plantations. Old plantations (65–72 years) of four different species, namely Araucaria cunninghamii, Agathis robusta, Toona ciliata and Flindersia brayleyana, as well as natural secondary forest were examined at Wongabel State Forest, in the wet tropics region of Queensland, Australia. Two young plantations (23 years) of Araucaria cunninghamii and Pinus caribaea were also examined. The close proximity of the older plantations and natural forests meant they had similar edaphic and climatic conditions. All plantations had been established as monocultures, but had been colonised by a range of native woody plants from the nearby rainforest. The extent to which this had occurred varied with the identity of the plantation species (from 2 to 17 species in 0.1 ha blocks). In many cases these additional species had grown up and joined the forest canopy. This study is one of the few to find a negative relationship between overstorey plant diversity and productivity. The conversion of natural forest with highly productive, low-diversity gymnosperm-dominated plantations (young and old Araucaria cunninghamii and Pinus caribaea) was found to be associated with lower soil nutrient availability (approximately five times less phosphorus and 2.5 times less nitrogen) and lower soil pH (mean = 6.28) compared to the other, less productive plantations. The dominant effects of two species, Araucaria cunninghamii and Hodgkinsonia frutescens, indicate that ecosystem functions such as production and nutrient availability are not determined solely by the number of species, but are more likely to be determined by the characteristics of the species present. This suggests that monoculture plantations can be used to successfully restore some functions (e.g. nutrient cycling and production), but that the level to which such functions can be restored will depend upon the species chosen and site conditions. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Disturbance processes and spatial patterns of two emergent conifers in New Caledonia

Abstract Araucaria laubenfelsii Corbasson and Araucaria montana Brongn et Gris are emergent conifers in maquis and forest communities which are subjected to a combination of fire and cyclonic disturbances. Both species are able to survive fire once stems reach a critical size, but most seedlings and saplings are killed. Both species were found to be clumped at most spatial scales for both saplings and trees in maquis, probably resulting from a combination of patchy fire and a limited ability to disperse seeds. Comparisons of burned and unburned A. montana sites suggest that clumping increases only slightly after fire. The degree of clumping in the A. laubenfelsii sites, suggested that fire disturbances may occur relatively frequently in the region. Cyclonic disturbances are infrequent but may result in blow‐down of large individuals within both maquis and forest. All of the individuals blown‐down during the study had been previously fire‐scarred. Around tree blow‐downs, seedling and sapling densities can be high. This likely reflects both the low dispersability of Araucaria seeds and enhanced moisture from the shading of the adult, when it was alive. Disturbance by fire and wind play an important role in the regeneration dynamics and spatial pattern of these species in maquis in New Caledonia.     

Investigation on cell death in the megagametophyte of Araucaria bidwillii Hook. post-germinated seeds

The megagametophyte of the Araucaria bidwillii seed is a storage tissue that surrounds and feeds the embryo. When all its reserves are mobilized, the megagametophyte degenerates as a no longer needed tissue. In this work we present a biochemical and a cytological characterization of the megagametophyte cell death. The TUNEL assay showed progressive DNA fragmentation throughout the post-germinative stages, while DNA electrophoretic analysis highlighted a smear as the predominant pattern of DNA degradation and internucleosomal DNA cleavage only for a minority of cells at late post-germinative stages. Cytological investigations at these stages detected profound changes in the size and morphology of the megagametophyte nuclei. By using in vitro assays, we were able to show a substantial increase in proteolytic activities, including caspase-like protease activities during the megagametophyte degeneration. Among the caspase-like enzymes, caspase 6- and 1-like proteases appeared to be the most active in the megagametophyte with a preference for acidic pH. On the basis of our results, we propose that the major pathway of cell death in the Araucaria bidwillii megagametophyte is necrosis; however, we do not exclude that some cells undergo developmental programmed cell death.     

Conditioning of the culture medium by suspension cells and formation of somatic proembryo in Araucaria angustifolia (coniferae)

Summary Cell masses of Araucaria angustifolia cultured in LP liquid medium showed different uptake and utilization patterns for different sugars. Fructose was slowly utilized by cell growth during the lag phase. After this lag period, fructose concentration decreased quickly, corresponding to the start of linear growth. Glucose rapidly decreased following fructose depletion. The level of extracellular proteins in the culture medium increased for the first 5 d after cell inoculation, during the transition from the lag phase to the high-growth phase. The pH of the medium decreased through the first half of the culture period (4.94) and increased (5.6) thereafter. Proembryos originated from a single cell, without a cleavage process. Suspensor cells and proembryonal groups developed independently. Only embryogenic cells divided and formed a suspensor or a proembryo group, which differentiated further along the longitudinal axis. The pathway observed for proembryo formation in Araucaria differs from the model proposed for other conifers, which show cleavage polyembryony.     

Desiccation studies in relation to the storage of Araucaria seed

Relationships between seed moisture content (fresh weight basis) and germination were examined for nine Araucaria species by desiccation under mild environmental conditions. The lowest safe moisture content, below which germination percentage begins to decline, was estimated in each case. Seeds can be grouped into three moisture content categories: the first group (including A. araucana, A. angustifolia, A. hunsteinii and A. bidwillii) cannot be safely dried to below 25–40%; the second group (including A. columnaris, A. rulei, A. nemorosa and A. scopulorum) cannot be dried to below about 12% without damage; the third category contains A. cunninghamii, which can be dried to 2% without damage. Seeds in the first group should be stored at 0–5 °C with moisture contents above the lowest-safe value. Provided freezing damage does not exceed 10%, seeds in the second group should be kept at - 18°C or lower with about 7% moisture content for long-term storage and at 0–5 °C with about 12% moisture content in the short term. Seed of A. cunninghamii is best retained at near 5% moisture content and in -18°C or lower. The lowest-safe moisture content was found to be associated with seed size and weight, higher moisture content values coinciding with greater size and weight of seed. Food reserve materials also differed among the groups; seeds of the first group were mainly starchy, whilst those in the other categories possessed a high lipid content.     

Indigenous Knowledge and Management of Araucaria Araucana Forest in the Chilean Andes: Implications for Native Forest Conservation

Southern Chile experienced serious deforestation during the past century and it is projected that by the year 2025 Chile will be devoid of native forests. One of the most important endemic tree species of the country and at the same time one of the most endangered ones is Araucaria araucana (Mol.) C. Koch, the monkey-puzzle tree. It grows in the Andes Mountains, homeland of the indigenous Mapuche Pewenche people who depend on this tree. This paper is based on participatory field research with a Mapuche Pewenche community in the southern Chilean Andes on their ecological knowledge, values, use and management of the Araucaria araucana forest. It attempts to reveal how indigenous people and their knowledge contribute to the sustainable management of these forests. The paper (1) illustrates the complexity of indigenous ecological knowledge of Araucaria araucana and its efficacy in native forest management, (2) explores the link between the conservation and use of biodiversity by the indigenous people, and (3) provides answers relevant to native forest management and conservation strategies ex-situ and in-situ incorporating indigenous and scientific knowledge, thus providing a contribution towards integrated natural resource management.     

Patterns of habitat utilization of small rodents in an area of Araucaria forest in Southern Brazil

Abstract We investigated patterns of spatial distribution and microhabitat selection of rodents in a patch (26 ha) of mixed ombrophilous forest (or Araucaria forest), dominated by the Araucaria ‘pine’Araucaria angustifolia, in the National Park of Aparados da Serra, RS, in southern Brazil. We conducted monthly trapping sessions lasting 6 days (from June 2000 to June 2001) using a square grid of 121 trap stations, separated by 15 m (2.25 ha). We measured 10 microhabitat variables that could influence species distribution. These variables were related to the occurrence of mammals by using canonical correspondence analysis (CCA) and stepwise multiple regression. The distribution patterns of the three most common rodents differed significantly (χ² = 38.6; P < 0.002), with Delomys dorsalis being more clumped than the other two species. Both CCA and multiple regression analysis showed a significant effect of microhabitat variables on species occurrence. The multiple regression parameters for Oligoryzomys nigripes were R² = 0.142, P < 0.05, for Akodon montensis were R² = 0.222, P < 0.01, and for D. dorsalis R² = 0.323, P < 0.001. Variables that significantly influenced species distribution were vegetation density at 1 m height and length of fallen logs for O. nigripes, canopy cover for A. montensis, and vegetation density at 2 m height, quantity of leaves of Araucaria trees (negative relationship), tree density (except Araucaria trees), and number of potential shelters, for D. dorsalis. The results indicated that D. dorsalis is the most habitat specialist among the rodent species and that there are no apparent competitive interactions.     

Grain protein variability among species of Triticum and Aegilops: quantitative SDS-PAGE studies

Summary Total proteins were extracted from degermed seeds of various species of Triticum and Aegilops with solutions containing sodium dodecyl sulfate (SDS) and mercaptoethanol. The reduced, dissociated proteins were fractionated according to molecular weight (MW) by high-resolution polyacrylamide gel electrophoresis in buffers containing SDS (SDS-PAGE). Stained SDS-PAGE patterns were measured by densitometric scanning over a suitable range of optical density. The data were normalized to equivalent total areas for each of the densitometric scans by means of a computer program that also permitted the construction of patterns of hypothetical amphiploids by averaging patterns of two or three diploid species. The grain proteins of most species examined had distinctive qualitative and quantitative aspects that were characteristic of the species even though nearly every accession or cultivar of a species exhibited at least minor differences in pattern from other accessions or cultivars. The main protein components (probably prolamins) of Triticum monococcum ssp. monococcum, T. monococcum ssp. boeoticum, T. urartu, and Aegilops squarrosa had MW's in the range 29–36 X 10³ whereas the most important components of Ae. speltoides, Ae. longissima, and Ae. searsii had MW's in the range 37–55 × 10³. Changes in the quantitative expression of particular genes, especially those coding for storage protein components, may have been associated with speciation. The strong predominance of proteins with MW's in the range 29–36 × 10³ in some accessions of AB genome tetraploids, such as T. turgidum ssp. dicoccoides, may indicate contributions to the B genome of these tetraploids by T. monococcum ssp. boeoticum, T. urartu, or Ae. squarrosa.     

Characterization of nuclear microsatellite loci in South American Araucariaceae species

The Araucariaceae family has only two species in South America: Araucaria angustifolia and Araucaria araucana. Both species are mainly used for timber and have been overexploited in the past. Currently, they are found as fragmented populations and are classified under the International Union for Conservation of Nature and Natural Resources (IUCN) guidelines as vulnerable species. Population fragmentation may seriously affect the genetic diversity of these two species of Araucariaceae and can consequently lead to decreased survival. To better understand the genetic structure of these South American Araucaria species, eight nuclear microsatellites are reported: six new microsatellites loci developed based on a membrane enrichment procedure and two microsatellites loci transferred from the related species, Araucaria cunninghamii.     

Storage protein dynamics in zygotic and somatic embryos of white fir

Composition and accumulation patterns of storage proteins in female gametophyte and embryos of the white fir (Abies concolor) were investigated during embryogenesis and germination of mature seeds using SDS-PAGE and immunological approach. Altogether 9 major and minor protein components with molecular masses of 14, 16, 22, 24, 27, 30, 35, 38, and 43 kDa were detected in female gametophytes and 9 protein bands in the embryos with the molecular sizes of 14, 16, 22, 24, 25, 27, 34, 38, and 43 kDa. The species seems to deviate in this respect from other representatives of Pinaceae. A conspicuous increase of storage protein synthesis was observed at the stage of fully cellularized female gametophytes and at the cotyledonary stage of embryo development. There exists a high degree of similarity between storage protein profiles of white fir zygotic and somatic embryos. Successive stages of somatic embryogenesis exhibited a high degree of similarity of storage proteins except for cotyledonary stage when a noticeable increase in storage protein synthesis was registered. Conversely, during germination of somatic embryos, an overwhelming majority of storage proteins was depleted.     

Ascorbate and glutathione metabolism in embryo axes and cotyledons of germinating lupine seeds

Changes in ascorbate and glutathione contents and the activities and isoenzyme patterns of enzymes of the ascorbate-glutathione cycle were investigated in embryo axes and cotyledons of germinating lupine (Lupinus luteus L.) seeds. Ascorbate content was not significantly affected over the initial 12 h of imbibition in embryo axes, but afterwards increased, with the most rapid accumulation coinciding with radicle emergence. A somewhat similar trend was observed for glutathione with significant increase in embryo axes shortly before radicle protrusion followed by decline in the next hours. In cotyledons the ascorbate pool rose gradually during germination but the amount of glutathione showed fluctuations during a whole germination period. The activity of ascorbate peroxidase (APX) rose progressively in embryo axes, while activities of dehydroascorbate reductase (DHAR) and glutathione reductase (GR) showed transient increase during germination. New isoforms of APX and GR were synthesized, suggesting that they play a relevant role during germination. All analyzed enzymes were already present in dry seeds which allowed them to be active immediately after imbibition.     

Proteins in development and germination of a desiccation sensitive (recalcitrant) seed species

In the recalcitrant seeds of Avicennia marina, protein content and the rates of protein synthesis increase during histodifferentiation. This is similar to the situation in desiccation tolerant seeds. During the stage of reserve accumulation the protein content and rates of synthesis remain constant and there is no de novo synthesis of proteins which might qualify as storage proteins. There is also no change in the nature of proteins present in either axis or cotyledonary tissues during development or germination. Similarly, fluorographs of axis proteins show only very limited changes in the patterns of protein synthesis during development and germination, at least until the onset of root growth. Heat-stable proteins are present from an early developmental stage. However, no late embryogenic abundant (LEA) proteins are synthesised during the late stages of development, indicating that seedling establishment is independent of such maturation proteins. It is suggested that the lack of desiccation tolerance of A. marina seeds might be related to the absence of desiccation-related LEAs. Although the rate of protein synthesis increases during germination, protein metabolism appears to remain qualitatively the same as that occurring during development. The present results suggest that in these desiccation sensitive seeds, protein metabolism characterising development changes imperceptibly into that of germination.     

The host range and biology of the mesquite psyllid Heteropsylla texana

Prosopis spp. areintroduced rangeland weeds in Australia. In asearch for biological control agents, thepsyllid Heteropsylla texana, which causessevere distortion to growing leaf and floralshoots, was imported from Texas, USA forhost range testing in quarantine facilities.No-choice tests were conducted on 60 plantspecies including P. pallida, P. velutinaand P. juliflora. In these trials, adultssurvived on 45 non-target plants but ovipositedonly on the Prosopis spp., Dichrostachys spicata and Acaciabidwillii. Nymphs developed to ovipositingadults on all Prosopis spp. and both D. spicata and A. bidwillii. Developmentof small numbers of adults on D. spicataand A. bidwillii occurred for only onegeneration. Damage was noticed only on Prosopis spp. In multiple-choice trials usingthree plant species, oviposition and subsequentdevelopment of eggs and nymphs to adults tookplace on P. pallida and D. spicatabut not on A. bidwillii. The low numbersdeveloping, the longer development times toadults when reared on D. spicata and A.bidwillii, and the failure of these plantsto sustain populations beyond one generationindicate that these plants are not hosts ofH. texana. While adult survival on manytest plants may imply that adult feedingoccurred, the risk to populations of theseplants in the field is negligible. It wasconcluded that H. texana is specific toProsopis spp. and could be released inAustralia for control of Prosopis spp.Observations of the biology of this speciesmade during the course of rearing andexperimentation indicated that nymphs developedthrough five instars to adults in 7–8 days sothat total development from egg to adult takes13–17 days. Females produce up to 100 eggs.     

Seed dispersal by the Florida box turtle (<Emphasis Type="Italic">Terrapene carolina bauri</Emphasis>) in pine rockland forests of the lower Florida Keys,United States

Seed dispersal by animals is one of the most important plant-animal mutualisms, but saurochory, the dispersal of seeds by reptiles, has received little attention. We investigated the role of the Florida box turtle (Terrapene carolina bauri) as a seed dispersal agent in pine rockland forests of the lower Florida Keys and examined the effect of turtle digestion on seed germination. We obtained seeds of 11 species with fleshy fruits and 2 species with non-fleshy fruits (a grass and legume) from the feces of 145 box turtles collected on Key Deer National Wildlife Refuge from 1999 to 2000. We planted the seeds of nine species and germination percentage (percentage of seeds that germinated during the experiment) varied from 10% to 80%. Comparative germination experiments were conducted with Thrinax morrissii, Serenoa repens, and Byrsonima lucida. We compared the germination percentage and germination rate (number of days from planting to seedling emergence) of seeds from three treatments (seeds recovered from feces, control seeds with pulp, and control seeds without pulp) and continued these experiments for up to 2 years. Passage through the box turtle digestive tract greatly enhanced the germination percentage and germination rate of S. repens, but decreased the germination percentage of B. lucida and T. morrissii, and decreased germination rate for T. morrissii. Subsequent destructive seed viability tests revealed that many ungerminated T. morrissii seeds remained viable, suggesting long-term seed dormancy may occur, even after passage through the turtle digestive system. In addition, the proportion of ungerminated seeds which remained viable was greater for seeds recovered from turtle feces than from control seeds with pulp. Furthermore, removal of fleshy pulp either manually or by the turtle digestive system may allow T. morrissii to escape insect predation.