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PubMed1999

Relationship between amount of esterase and gene copy number in insecticide-resistant Myzus persicae (Sulzer).

Overproduction of the insecticide-degrading esterases, E4 and FE4, in peach-potato aphids, Myzus persicae (Sulzer), depends on both gene amplification and transcriptional control, the latter being associated with changes in DNA methylation. The structure and function of the aphid esterase genes have been studied but the determination of their copy number has proved difficult, a common problem with gene amplification. We have now used a combination of pulsed-field gel electrophoresis and quantitative competitive PCR to determine relative esterase gene copy numbers in aphid clones with different levels of insecticide resistance (R1, R2 and R3). There are approx. 4-fold increases between susceptible, R1, R2 and R3 aphids, reaching a maximum of approx. 80 times more genes in R3; this gives proportionate increases in esterase protein relative to susceptible aphids. Thus there is no overexpression of the amplified genes, in contrast with what was thought previously. For E4 genes, the loss of 5-methylcytosine is correlated with a loss of expression, greatly decreasing the amount of enzyme relative to the copy number.

5-Methylcytosine↗
PubMed2001

[Effects of volatiles from different trophic level on foraging behavior of Aphidius avenae].

The responses of female Aphidius avenae to volatiles from undamaged wheat plant, Sitobion avenae and Rhopalosiphum padi, aphids-plant complexes and aphids damaged plant were investigated in wind tunnel and Y-tube olfactometer. The volatiles from undamaged wheat plant, S. avenae and R. padi were much less attractive to A. avenae than those from aphids-plant complexes and aphids-damaged plant. Although the parasitic rate was much lower, the R. padi-plant complexes and R. padi damaged plant were also attractive to A. avenae. Identification of volatiles by GC-MS showed that the main volatiles induced by aphids feeding were 2-camphene, 6-methyl-5-hepten-2-one, 6-methyl-5-hepten-2-ol, Z-3-hexenyl-acetate, and methyl salicylate, among which, 6-methyl-5-hepten-2-one and 6-methyl-5-hepten-2-ol were more attractive to A. avenae, while methyl salicylate was not attractive.

Animals↗
PubMed2003

Effectiveness of predators and parasitoids for the natural control of Diuraphis noxia (Homoptera: Aphididae) on barley in central Ethiopia.

The Russian wheat aphid, Diuraphis noxia (Kurdj.), is a serious threat for the production of barley, Hordeum vulgare L., in Ethiopia. The pest can cause up to 100% yield losses in years of heavy infestation. Natural control of D. noxia by using predators and parasitoids might be the best alternative both from economical and ecological point of view. In this regard, a field investigation was conducted during the 2001 spring cropping season at Fitche, central Ethiopia to determine abundance of D. noxia and associated natural enemies and evaluate their effectiveness as natural control agents for this pest. In addition, the data collected during the 1995-1997 spring and summer cropping seasons at Chacha was analysed and presented. There was a highly significant difference (P < 0.01) on the population density of D. noxia between spring and summer and among sampling dates. The average aphid density was 23.1 and 0.7 aphids per tiller during spring and summer, respectively. The quantitative analyses based on predator units (PU) showed a highly significant difference (P < 0.01) among the predator groups, coccinellids and syrphids being more important, however, the coccinellid beetle, Hippodamia variegata Goeze, was the dominant species from all predator species or groups. Moreover, syrphid fly larvae were the only predators consistently found feeding on aphids within D. noxia rolled leaves. The polyphagous predators: Chrysopids, spiders and staphylinids were found to have less effect on D. noxia populations. Only two primary parasitoid species were recorded both during spring and summer cropping seasons: Aphidius colemani Vier. and Aphidius spp. Of these, A. colemani was more abundant in both seasons. The rate of parasitism was also generally low during the season. In general, the population build up of these natural enemies is slow and maximum population size is reached after the aphid population peaks and the crop has already been damaged. Further extensive work is necessary to maximize the effectiveness of natural enemies against this pest.

Animals↗
PubMed2005

The use of common elder Sambucus nigra to promote Aphidophagous syrphids in apple orchards.

Elder shrubs (Sambucus nigra L.) were planted in an experimental apple orchard as bordering hedgerow with the objective of rearing syrphids (Diptera: Syrphidae) and consequently enhancing the biological control of the rosy apple aphid Dysaphis plantaginea (Passerini). The inoculative introduction of the specific elder aphid Aphis sambuci L. (Homoptera: Aphididae) in late March 2002 and again in 2003 led to the complete hedgerow turning into an early and productive alternative prey reservoir for indigenous syrphids. The species Scaeva pyrastri (L.), S. selenetica (Meigen), Syrphus ribesii (L.), S. vitripennis Meigen, Epistrophe eligans (Harris), E. nitidicollis (Meigen), Platycheirus scutatus (Meigen), Eupeodes corollae (F.), Meligramma triangulifera (Zetterstedt) and Episyrphus balteatus (De Geer) developed on the elder hedgerow during the spring 2002 taking advantage of the aphid infestation. The season 2003 was characterised by a quite different species abundance pattern as, contrary to the previous year when both monovoltine and polyvoltine species were equally represented, the monovoltine syrphids (Epistrophe spp.) represented the dominant group exploiting the elder aphids. A parallel analysis showed that the faeces of the first adult syrphids observed ovipositing on the elder hedgerow in spring contained digested pollen of mainly apple. Our observations indicate the possibility of establishing a local population of monovoltine syrphids in apple orchards by managing an aphid-infested elder hedgerow, without any additional pollen or nectar producing plants.

Animals↗
PubMed2002

Intraguild interference and biocontrol effects of generalist predators in a winter wheat field.

Arable land typically harbours communities of polyphagous invertebrate natural enemies, among them numerous soil-surface dwelling predators such as ground beetles (Carabidae) and spiders (Lycosidae, Linyphiidae). Numbers of these predators were experimentally manipulated in a winter wheat field in order to study the predation impact of a generalist predator assemblage on herbivorous insects, the possible interferences among the predators concerned, and subsequent effects on wheat plant parameters. Removing ground beetles doubled numbers of Lycosidae indicative of intraguild interference between these two predator groups. Aphid densities were highest in carabid removal plots implying a substantial predation impact of ground beetles on the pest population. The predation impact of ground beetles was strongest earlier and disappeared later in the season. In mid-season, at intermediate aphid densities, the combined impact of carabid beetles and spiders appeared to be responsible for the reduction in aphid abundance. This result was probably due to a biomass effect rather than to a synergistic effect of the predator community. Thysanoptera decreased when spiders were removed (perhaps because spiders were preying on a predator of thrips), while Cicadellidae and Delphacidae showed no effect at all. The rise of aphid numbers in carabid removal plots corresponded to an increase in protein content of the wheat grains, while other plant parameters such as plant numbers and grain mass were not affected. In conclusion, this study provided field evidence for intraguild interference among generalist ground predators in arable land. Despite this interference the polyphagous predator community was able to depress numbers of aphids in winter wheat, a result cascading down to plant quality parameters.

Animals↗
PubMed2004

Suppression of insect pest populations and damage to plants by vermicomposts.

The effects of commercial vermicomposts, produced from food waste, on infestations and damage by aphids, mealy bugs and cabbage white caterpillars were studied in the greenhouse. Vermicomposts were used at substitution rates into a soil-less plant growth medium, MetroMix 360 (MM360), at rates of 100% MM360 and 0% vermicompost, 80% MM360 and 20% vermicompost, and 60% MM360 and 40% vermicompost to grow peppers (Capsicum annuum L.), tomatoes (Lycopersicon esculentum Mill.) and cabbages (Brassica oleracea L.), in pots. Groups of 10 pots containing young plants were distributed randomly in nylon mesh cages (40 cm x 40 cm x 40 cm). Groups of 10 pepper seedlings in a single cage were infested with either 100 aphids (Myzus persicae Sulz.) or 50 mealy bugs (Pseudococcus spp.) per cage. Similar groups of tomato seedlings were infested with 50 mealy bugs per cage. Groups of four cabbage seedlings in pots in cages were infested with 16 cabbage white caterpillars (Pieris brassicae L.). Populations of aphids and mealy bugs were counted after 20 days and the shoot dry weights of peppers, tomatoes and cabbages were measured at harvest. Numbers of cabbage white caterpillars and loss in shoot weights were measured after 15 days. The substitution rates of 20% and 40% vermicomposts suppressed populations of both aphids and mealy bugs on peppers, and mealy bugs on tomatoes, significantly. Substitutions with vermicomposts into MM360 decreased losses of dry weights of peppers, in response to both aphid and mealy bug infestations, decreased losses in shoot dry weights of tomatoes after mealy bug infestations significantly. There were significantly decreased losses in leaf areas of cabbage seedlings in response to the cabbage white caterpillar infestations.

Analysis of Variance↗
PubMed1998

Host regulation effects of ovary fluid and venom of Aphidius ervi (Hymenoptera: Braconidae).

The host regulation effects of venom and ovary fluid of the endophagous braconid Aphidius ervi Haliday on the pea aphid, Acyrthosiphon pisum (Harris), have been studied. Extracts of ovaries and of venom glands were injected into nonparasitized 4th instar pea aphids, both separately and mixed. Aphids treated with parasitoid material died as 4th instars, often showing developmental arrest. In contrast, control aphids that received an injection of Pringle's saline solution regularly moulted to the adult stage and reproduced. Venom alone was as effective as the combined extracts in determining developmental arrest and death. Separate heat and protease treatments of these parasitoid's reproductive secretions significantly reduced their biological activity, suggesting that the active component(s) involved is a protein(s). SDS-PAGE analysis of haemolymph samples obtained from pea aphids which had received an injection of combined venom and ovary extract revealed an increase of the titre of various proteins, particularly in the 43-47kDa interval, as registered for truly parasitized hosts. This altered protein profile was first detected 48h following injection. Based on this information a tentative physiological model is proposed. The apical tract of host ovarioles, where the germarium and growing oocytes are located, is suggested to be one of the major targets of female parasitoid secretions injected with the egg.

Journal Article↗
PubMed1998

Use of modified plum pox virus coat protein genes developed to limit heteroencapsidation-associated risks in transgenic plants.

Aphid transmission of a non-aphid-transmissible strain of zucchini yellow mosaic virus (ZYMV-NAT) occurs in transgenic plants expressing the plum pox potyvirus (PPV) coat protein (CP) gene. Heteroencapsidation has been shown to be responsible for this modification in the epidemiological characteristics of the infecting virus. In order to prevent this biological risk, several modified PPV CP constructs were produced that were designed to interfere with heteroencapsidation itself or to block aphid transmission of heteroencapsidated virions. These constructs were first expressed in Escherichia coli in order to check for the accumulation of pseudo-particles by electron microscopy. Virus-like particles (VLPs) were found with the full-length CP and with a PPV CP lacking the DAG amino acid triplet involved in aphid transmission. However, no VLPs were observed with CP lacking R220, Q221 or D264, amino acids known to be essential for the assembly of other potyvirus CPs. Transgenic Nicotiana benthamiana lines expressing the different PPV CP constructs were infected with ZYMV-NAT. Aphid transmission assays performed with these plants demonstrated that the strategies developed here provide an effective means of minimizing the biological risks associated with heteroencapsidation.

Capsid↗
PubMed2005

Insecticidal activity of 23 essential oils and their major compounds against adult Lipaphis pseudobrassicae (Davis) (Aphididae: Homoptera).

Essential oils from 23 species of plants comprising 14 genera and 4 plant families were obtained by Clevenger-type water distillation. The major compounds in these essential oils were identified with GC-MS and their insecticidal activity against adult turnip aphids, Lipaphis pseudobrassicae (Davis), tested with dosage-mortality bioassays. We examined mortality only for viviparous adults because sizeable aphid populations on crucifer (Brassicaceae) hosts are largely produced by these wingless, parthenogenic females. Twenty-two of the oils were directly applied to aphid females in randomized blocks at concentrations of 0.0, 1.0, 2.5, 5.0 and 10.0 mg ml(-1). Essential oils mixed with a non-toxic emulsifying agent, dimethyl sulfoxide (DMSO), more easily penetrated the waxy insect cuticle. Probit analysis and LC(50) at three different exposures showed aphids were quickly incapacitated and killed by aliphatic aldehydes, phenols and monocyclic terpenes contained in Bifora and Satureja oils and at applied concentrations as low as 0.3 to 1.0 mg ml(-1). Only enough Pimpinella isaurica oil and its three phenylpropanoid fractions were available for testing at a single concentration of 10 mg ml(-1). We could not spare any additional P. isaurica oil for testing at other concentrations. Phenylpropanoids isolated from P. isaurica oil when recombined or left naturally blended in the oil were highly bioactive against L. pseudobrassicae at 10 mg ml(-1).

Aging↗
PubMed2005

Development and reproduction of Adalia bipunctata (Coleoptera: Coccinellidae) on eggs of Ephestia kuehniella (Lepidoptera: Phycitidae) and pollen.

Due to growing criticism over the use of non-indigenous coccinellids, the two-spot ladybird, Adalia bipunctata (L.), has enjoyed increasing attention for aphid biocontrol in Europe. In the current study, eggs of the Mediterranean flour moth, Ephestia kuehniella Zeller, whether or not supplemented with bee pollen, were evaluated as a factitious food for larvae and adults of A. bipunctata. The predator showed slower larval development and lower survival when reared on live pea aphids, Acyrthosiphon pisum (Harris), than on E. kuehniella eggs. Survival on gamma-irradiated eggs of E. kuehniella was superior to that on frozen flour moth eggs, but other developmental characteristics were similar. Adults of A. bipunctata reared on Ac. pisum were only half as fecund as those offered irradiated or frozen E. kuehniella eggs, but egg hatch was markedly better on live aphids than on flour moth eggs (61 versus 20-27%, respectively). However, when a diet of flour moth eggs was supplemented with frozen moist bee pollen, egg hatch of A. bipunctata was equally as good as on live aphids. Supplementing flour moth eggs with dry pollen did not yield satisfactory results. Only 10% of larvae reached adulthood on moist bee pollen alone and resulting adults weighed less than half as much as those obtained on flour moth eggs. Our findings indicate that A. bipunctata is able to compensate for a suboptimal diet of animal prey by supplementary feeding on flower pollen. It is concluded that pollinivory may be a crucial trait for both the rearing of this natural enemy and its use in biological control programmes.

Animals↗
PubMed2006

Differential aphicidal effects of chitinase inhibitors on the polyphagous homopteran Myzus persicae (Sulzer).

Four chitinase inhibitors, cyclo-(Proline-Tyrosine), cyclo-(Histidine-Proline), allosamidin and psammaplin A, were selected for in vitro feeding experiments with the peach-potato aphid, Myzus persicae (Sulzer), under controlled photoperiod and temperature conditions. Artificial diets were used to provide chitinase inhibitors at 10, 50 and 100 microg mL(-1) to M. persicae. Except for cyclo-(Proline-Tyrosine), which did not modify aphid demographic parameters, chitinase inhibitors induced differential aphicidal effects on M. persicae. At all doses, cyclo-(Histidine-Proline) induced significant effects affecting daily fecundity, intrinsic rate of natural increase (r(m)) and doubling time of population. When compared with the control diet, allosamidin decreased nymph survival and daily fecundity, increasing the doubling time of population from 1 to 1.5 days. Psammaplin A was the most toxic inhibitor when delivered via artificial diet, as it induced the death of all aphids reared at 50 and 100 microg mL(-1). The results demonstrate the potential use of chitinase inhibitors as aphid management tools.

Acetylglucosamine↗
PubMed2004

Occurrence of barley yellow dwarf virus in autumn-sown cereal crops in the United Kingdom in relation to field characteristics.

The incidence of barley yellow dwarf virus and of its aphid vectors was surveyed in untreated parts of autumn-sown cereal crops, mainly wheat and barley, in the United Kingdom in 1995-8. The incidence of virus in the spring was related to the incidence of aphids in the preceding autumn. Both virus and aphid incidences could also be related to a range of crop and field characteristics, in particular sowing date, regions as defined by their geographical position, topography and climate, the proximity of the field to the sea, the extent of arable land in the vicinity of the field, and the aspect and size of the field. Proximity of cropped fields to setaside areas had no significant effect on either aphid or virus incidence.

Animals↗
PubMed2001

Parallel acceleration of evolutionary rates in symbiont genes underlying host nutrition.

The overproduction of essential amino acids by Buchnera aphidicola, the primary bacterial mutualist of aphids, is considered an adaptation for increased production of nutrients that are lacking in aphids' diet of plant sap. Given their shared role in host nutrition, amino acid biosynthetic genes of Buchnera are expected to experience parallel changes in selection that depend on host diet quality, growth rate, and population structure. This study evaluates the hypothesis of parallel selection across biosynthetic pathways by testing for correlated changes in evolutionary rates at biosynthetic genes of Buchnera. Previous studies show fast evolutionary rates at tryptophan biosynthetic genes among Buchnera associated with the aphid genus Uroleucon and suggest reduced purifying selection on symbiont nutritional functions in this aphid group. Here, we test for parallel rate acceleration at other amino acid biosynthetic genes of Buchnera-Uroleucon, including those for leucine (leuABC) and isoleucine/valine biosynthesis (ilvC). Ratios of nonsynonymous to synonymous substitutions (d(N)/d(S)) were estimated using codon-based maximum-likelihood methods that account for the extreme AT compositional bias of Buchnera sequences. A significant elevation in d(N)/d(S) at biosynthetic loci but not at two housekeeping genes sampled (dnaN and tuf) suggests reduced host-level selection on biosynthetic capabilities of Buchnera-Uroleucon. In addition, the discovery of trpEG pseudogenes in Buchnera-U. obscurum further supports reduced selection on amino acid biosynthesis.

Alcohol Oxidoreductases↗
PubMed2005

Induced plant responses to multiple damagers: differential effects on an herbivore and its parasitoid.

Herbivore-induced plants responses can affect the preference and performance of herbivores and their natural enemies. These responses may vary depending on the identity and number of herbivore species feeding on the plant so that when herbivores from different guilds feed on plants, the interactions between plants, herbivores, and natural enemies may be disrupted. Tomato plants were damaged either by the caterpillar Spodoptera exigua, or the aphid Macrosiphum euphorbiae, or damaged by both herbivores, or undamaged controls. We measured the preference and performance of S. exigua and its parasitoid Cotesia marginiventris, and activity of proteinase inhibitors (PI) as an indicator of induced resistance. Compared to undamaged plants, caterpillar damage reduced the number of eggs laid by S. exigua adults, reduced growth, consumption, and survival of larval S. exigua and C. marginiventris, and increased activity of PIs 43%; but did not increase attraction of C. marginiventris. While pupal mass of S. exigua was not affected, the pupal mass of C. marginiventris decreased on caterpillar-damaged plants compared to controls. In contrast, plants damaged by aphids were preferred for oviposition by S. exigua, and had increased larval consumption and survival, compared to controls. Aphid feeding did not affect the preference or performance of C. marginiventris, or PI activity, compared to controls. While oviposition was deterred on caterpillar-damaged plants, plants damaged by both herbivores received the same amount of oviposition as controls. The attraction of C. marginiventris to plants damaged by caterpillars and aphids was increased compared to controls. However, plants damaged by both herbivores had similar PI activity, larval growth and survival of S. exigua and C. marginiventris, as plants singly damaged by caterpillars. Overall, the preference component for both the herbivore and parasitoid was more strongly affected by damage due to multiple herbivores than the performance component.

Analysis of Variance↗
PubMed2005

The effects of ants on the entomophagous butterfly caterpillar Feniseca tarquinius, and the putative role of chemical camouflage in the Feniseca-ant interaction.

Butterfly caterpillars in the lycaenid subfamily Miletinae are predators of ant-tended Homoptera, yet they lack specialized secretory and call-production organs crucial to ant association in other lycaenids. Here, we address the question of how miletine caterpillars have invaded the ant-Homoptera symbiosis through a study of the only New World miletine, Feniseca tarquinius, a predator of the wooly aphid Prociphilus tesselatus. Previous interpretations have suggested that F. tarquinius and other miletine caterpillars avoid ant aggression by concealing themselves under silken webs. In contrast, our field data indicate that F. tarquinius caterpillars are less likely to be concealed in the presence of the ants Camponotus pennsylvanicus and Formica obscuriventris than in the absence of ants, although caterpillar and ant behaviors vary between years. Chemical analysis and behavioral assays suggest that chemical camouflage, not physical concealment, is responsible for the ants' failure to detect and remove F. tarquinius caterpillars from aphid colonies. Analyses by gas chromatography indicate that the cuticular lipid composition of caterpillars are similar to that of their aphid prey, although it varies with prey species. Behavioral assays confirm that solvent extracts of F. tarquinius caterpillars and P. tesselatus aphids evoke similar behavioral responses in C. pennsylvanicus ants. Chemical camouflage is well known in social parasites of ants, but the present study represents one of a few documented cases where chemical deceit is important to interactions with ants outside the nest.

Animals↗
PubMed1992

Site-directed mutations in the potyvirus HC-Pro gene affect helper component activity, virus accumulation, and symptom expression in infected tobacco plants.

Helper component (HC-Pro) is a virus-encoded nonstructural protein required for aphid transmission of potyviruses. In the tobacco vein mottling virus (TVMV) polyprotein, HC-Pro represents a 457 residue polypeptide from amino acid position 257 to 713. Previous sequence comparison studies have suggested that mutations of one or two specific amino acid residues in the HC-Pro protein might result in loss of aphid transmission activity. To test this hypothesis, the initial targets were the residues corresponding to these specific amino acids, a lys to glu change and an ile to val change at amino acid positions 307 and 482, respectively, of the TVMV polyprotein, as well as the combination of the two. Two additional mutations within the HC-Pro representing dipeptide changes thr-ser to ile-asp and thr-ala to leu-glu at amino acid positions (283/284) and (368/369), respectively, were also tested to further define the effects of mutations in this region on helper component activity. The mutations at positions 482 and (368/369) had no effect on aphid transmission activity, while mutation at position 307 completely abolished the activity. Except for the 482 mutation, all the mutations also affected symptomatology and virus accumulation in infected plants. Due to the very low concentrations of HC-Pro in plants infected with the (283/284) mutant, the effect of this dipeptide change on aphid transmission activity could not be assessed. The majority of the tested mutations fall within a putative zinc-finger motif postulated in the cysteine-rich N-terminus of HC-Pro. The possible role of this motif in the potyviruses is further discussed in the light of our present results with TVMV.

Animals↗
PubMed2006

Dispersal of Beauveria bassiana by the activity of nettle insects.

Recent studies have shown that the entomopathogenic fungus Beauveria bassiana occurs naturally on the phylloplanes of several plants, including nettles. Insects could, by their activity, be contributing to this inoculum by dispersing it from other sites. The potential of nettle aphids Microlophium carnosum and their predator Anthocoris nemorum to disperse conidia of B. bassiana from soil to nettles and from sporulating cadavers in the nettle canopy was investigated in laboratory experiments. In petri dish assays, aphids showed potential to distribute B. bassiana from soil to nettle leaves. Predators dispersed inoculum from both soil and cadavers to nettle leaves in petri dishes. In microcosms, aphids did not disperse B. bassiana from the soil or from cadavers confined in the canopy, but A. nemorum were able to transfer inoculum from soil into the nettle canopy and to distribute conidia from cryptic cadavers. In some instances, infections were initiated in aphids and predators as a consequence of dispersal.

Animals↗
PubMed2004

Speciation by host-switching in pinyon Cinara (Insecta: Hemiptera: Aphididae).

Parasite-host cospeciation has received much attention as an important mechanism in the diversification of phytophagous insects. However, studies have shown that for certain taxa, it is not host fidelity but host-switching that plays the critical role in speciation. Cinara are aphids (Insecta: Hemiptera: Aphididae: Lachninae) that feed exclusively on the woody parts of conifers of the Cupressaceae and Pinaceae. They are unusual aphids because most Pinaceae play host to several species of Cinara. The aphids show relatively strong host fidelity, and as a consequence historically have been treated based on the taxonomy of their hosts. The historical paradigm of aphid evolution implies that Cinara species have radiated to different parts of the same host species and/or speciated with their host. Using mitochondrial cytochrome oxidase 1 and nuclear elongation factor 1-alpha DNA sequences, we performed molecular phylogenetic analysis of Cinara species, concentrating on those associated with pinyon pines in the southwestern USA. We determined that switching hosts has played a key role in the speciation of the genus, reflected in the polyphyly of pinyon-feeding Cinara. Furthermore, species sharing a common feeding site on different hosts were more closely related to each other than to those sharing the same host but at different feeding sites, suggesting that feeding site fidelity plays a more important role in speciation than does host fidelity in general. This study also elucidated the primary taxonomy of various species: it suggested that Cinara rustica Hottes is a junior synonym of C. edulis (Wilson) and that C. wahtolca Hottes represents two species on the two different pinyon pine species, Pinus edulis Englem. and P. monophylla Torr. & Frem.

Animals↗