Biology Commons^™

Intramolecular Phenotypic Capacitance In A Modular Rna Molecule, Eric J. Hayden, Devin P. Bendixsen, Andreas Wagner

Biology Faculty Publications and Presentations

Phenotypic capacitance refers to the ability of a genome to accumulate mutations that are conditionally hidden and only reveal phenotype-altering effects after certain environmental or genetic changes. Capacitance has important implications for the evolution of novel forms and functions, but experimentally studied mechanisms behind capacitance are mostly limited to complex, multicomponent systems often involving several interacting protein molecules. Here we demonstrate phenotypic capacitance within a much simpler system, an individual RNA molecule with catalytic activity (ribozyme). This naturally occurring RNA molecule has a modular structure, where a scaffold module acts as an intramolecular chaperone that facilitates folding of a second …

An Irrigation Canal As A Lotic Mesocosm: Examining The Relationship Between Macroinvertebrate Benthos And Drift, Peter Koetsier, Luana M. M. Mccauley

Biology Faculty Publications and Presentations

We explored the notion that small canals could be good experimental proxies of streams by documenting physiochemical parameters and macroinvertebrate community development in an aridland irrigation canal. Further, we tested the production-compensation hypothesis between benthic invertebrates and invertebrates in the water column (drift). If the hypothesis held, invertebrates in the drift would be low until the benthic carrying capacity was reached; then organisms in the drift would increase as individuals avoided overcrowded conditions in the substrate. In a small, naturalized freshwater canal, we sampled macroinvertebrates in the substrate and those in the drift once every 2 weeks over 170 days …

A Phantom Road Experiment Reveals Traffic Noise Is An Invisible Source Of Habitat Degradation, Heidi E. Ware, Christopher J.W. Mcclure, Jay D. Carlisle, Jesse R. Barber

Biology Faculty Publications and Presentations

Decades of research demonstrate that roads impact wildlife and suggest traffic noise as a primary cause of population declines near roads. We created a “phantom road” using an array of speakers to apply traffic noise to a roadless landscape, directly testing the effect of noise alone on an entire songbird community during autumn migration. Thirty-one percent of the bird community avoided the phantom road. For individuals that stayed despite the noise, overall body condition decreased by a full SD and some species showed a change in ability to gain body condition when exposed to traffic noise during migratory stopover. We …

Caspase-Cleaved Tau Co-Localizes With Early Tangle Markers In The Human Vascular Dementia Brain, Ryan J. Day, Maria J. Mason, Chloe Thomas, Wayne W. Poon, Troy T. Rohn

Biology Faculty Publications and Presentations

Vascular dementia (VaD) is the second most common form of dementia in the United States and is characterized as a cerebral vessel vascular disease that leads to ischemic episodes. Whereas the relationship between caspase-cleaved tau and neurofibrillary tangles (NFTs) in Alzheimer’s disease (AD) has been previously described, whether caspase activation and cleavage of tau occurs in VaD is presently unknown. To investigate a potential role for caspase-cleaved tau in VaD, we analyzed seven confirmed cases of VaD by immunohistochemistry utilizing a well-characterized antibody that specifically detects caspase-cleaved tau truncated at Asp⁴²¹. Application of this antibody (TauC3) revealed consistent …

Origin And Evolution Of Petrocosmea (Gesneriaceae) Inferred From Both Dna Sequence And Novel Findings In Morphology With A Test Of Morphology-Based Hypotheses, Zhi-Jing Qiu, Yuan-Xue Lu, Chao-Qun Li, Yang Dong, James F. Smith, Yin-Zheng Wang

Biology Faculty Publications and Presentations

Background: Petrocosmea Oliver (Gesneriaceae) currently comprises 38 species with four non-nominate varieties, nearly all of which have been described solely from herbarium specimens. However, the dried specimens have obscured the full range of extremely diverse morphological variation that exists in the genus and has resulted in a poor subgeneric classification system that does not reflect the evolutionary history of this group. It is important to develop innovative methods to find new morphological traits and reexamine and reevaluate the traditionally used morphological data based on new hypothesis. In addition, Petrocosmea is a mid-sized genus but exhibits extreme diverse floral variants. This …

Pavement And Riparian Forest Shape The Bird Community Along An Urban River Corridor, Christopher J.W. Mcclure, Allison C. Korte, Julie A. Heath, Jesse R. Barber

Biology Faculty Publications and Presentations

Knowledge of habitat use by animals within urban-riparian corridors during the breeding season is important for conservation, yet remains understudied. We examined the bird community along an urban-riparian corridor through metropolitan Boise, Idaho and predicted that occupancy of individual species and species richness would be greater in forested areas than in urbanized areas. We surveyed birds throughout the summers of 2009 and 2010 and quantified the m² of each cover-type within 50-m, 100-m, and 200-m buffers surrounding each survey location using satellite imagery. Occupancy modeling revealed that eight of 14 species analyzed were positively associated with riparian forest, and …

Selfish Mitochondrial Dna Proliferates And Diversifies In Small, But Not Large, Experimental Populations Of Caenorhabditis Briggsae, Wendy S. Phillips, Anna Luella Coleman-Hulbert, Emily S. Weiss, Dana K. Howe, Sita Ping, Riana I. Wernick, Suzanne Estes, Dee R. Denver

Biology Faculty Publications and Presentations

Evolutionary interactions across levels of biological organization contribute to a variety of fundamental processes including genome evolution, reproductive mode transitions, species diversification, and extinction. Evolutionary theory predicts that so-called “selfish” genetic elements will proliferate when the host effective population size (N_e) is small, but direct tests of this prediction remain few. We analyzed the evolutionary dynamics of deletion-containing mitochondrial DNA (ΔmtDNA) molecules, previously characterized as selfish elements, in six different natural strains of the nematode Caenorhabditis briggsae allowed to undergo experimental evolution in a range of population sizes (N = 1, 10, 100, and 1,000) for a maximum …

Physiological Strategies During Animal Diapause: Lessons From Brine Shrimp And Annual Killifish., Jason E. Podrabsky, Steven C. Hand

Biology Faculty Publications and Presentations

Diapause is a programmed state of developmental arrest that typically occurs as part of the natural developmental progression of organisms that inhabit seasonal environments. The brine shrimp Artemia franciscana and annual killifish Austrofundulus limnaeus share strikingly similar life histories that include embryonic diapause as a means to synchronize the growth and reproduction phases of their life history to favorable environmental conditions. In both species, respiration rate is severely depressed during diapause and thus alterations in mitochondrial physiology are a key component of the suite of characters associated with cessation of development. Here, we use these two species to illustrate the …

Recbcd Is Required To Complete Chromosomal Replication: Implications For Double- Strand Break Frequencies And Repair Mechanisms, Justin Courcelle, Brian M. Wendel, Dena D. Livingstone, Charmain T. Courcelle

Biology Faculty Publications and Presentations

Several aspects of the mechanism of homologous double strand break repair remain unclear. Although intensive efforts have focused on how recombination reactions initiate, far less is known about the molecular events that follow. Based upon biochemical studies, current models propose that RecBCD processes double strand ends and loads RecA to initiate recombinational repair. However, recent studies have shown that RecBCD plays a critical role in completing replication events on the chromosome through a mechanism that does not involve RecA or recombination. Here, we examine several studies, both early and recent, that suggest RecBCD also operates late in the recombination process- …

Antioxidant Capacity Of Wyoming Big Sagebrush (Artemisia Tridentata Ssp. Wyomingensis) Varies Spatially And Is Not Related To The Presence Of A Sagebrush Dietary Specialist, Xinzhu Pu, Lisa Lam, Kristina Gehlken, Amy C. Ulappa, Janet Rachlow, Jennifer Sorensen Forbey

Biology Faculty Publications and Presentations

Sagebrush (Artemisia spp.) in North America is an abundant native plant species that is ecologically and evolutionarily adapted to have a diverse array of biologically active chemicals. Several of these chemicals, specifically polyphenols, have antioxidant activity that may act as biomarkers of biotic or abiotic stress. This study investigated the spatial variation of antioxidant capacity, as well as the relationship between a mammalian herbivore and antioxidant capacity in Wyoming big sagebrush (Artemisia tridentata wyomingensis). We quantified and compared total polyphenols and antioxidant capacity of leaf extracts from sagebrush plants from different spatial scales and at different levels …

A Meta-Analysis Of Soil Biodiversity Impacts On The Carbon Cycle, M.-A. De Graaff, J. Adkins, P. Kardol, H. L. Throop

Biology Faculty Publications and Presentations

Loss of biodiversity impacts ecosystem functions, such as carbon (C) cycling. Soils are the largest terrestrial C reservoir, containing more C globally than the biotic and atmospheric pools together. As such, soil C cycling, and the processes controlling it, has the potential to affect atmospheric CO₂ concentrations and subsequent climate change. Despite the growing evidence of links between plant diversity and soil C cycling, there is a dearth of information on whether similar relationships exist between soil biodiversity and C cycling. This knowledge gap occurs even though there has been increased recognition that soil communities display high levels of …

Cyclosporin A Disrupts Notch Signaling And Vascular Lumen Maintenance, Raghav Pandey, Mark A. Botros, Benjamin A. Nacev, Allan R. Albig

Biology Faculty Publications and Presentations

Cyclosporin A (CSA) suppresses immune function by blocking the cyclophilin A and calcineurin/NFAT signaling pathways. In addition to immunosuppression, CSA has also been shown to have a wide range of effects in the cardiovascular system including disruption of heart valve development, smooth muscle cell proliferation, and angiogenesis inhibition. Circumstantial evidence has suggested that CSA might control Notch signaling which is also a potent regulator of cardiovascular function. Therefore, the goal of this project was to determine if CSA controls Notch and to dissect the molecular mechanism(s) by which CSA impacts cardiovascular homeostasis. We found that CSA blocked JAG1, but not …

Oncostatin M Binds To Extracellular Matrix In A Bioactive Conformation: Implications For Inflammation And Metastasis, Randall E. Ryan, Bryan Martin, Liliana Mellor, Reed B. Jacob, Ken Tawara, Owen M. Mcdougal, Julia Thom Oxford, Cheryl L. Jorcyk

Biology Faculty Publications and Presentations

Oncostatin M (OSM) is an interleukin-6-like inflammatory cytokine reported to play a role in a number of pathological processes including cancer. Full-length OSM is expressed as a 26 kDa protein that can be proteolytically processed into 24 kDa and 22 kDa forms via removal of C-terminal peptides. In this study, we examined both the ability of OSM to bind to the extracellular matrix (ECM) and the activity of immobilized OSM on human breast carcinoma cells. OSM was observed to bind to ECM proteins collagen types I and XI, laminin, and fibronectin in a pH-dependent fashion, suggesting a role for electrostatic …

Mitigation Effectiveness For Improving Nesting Success Of Greater Sage-Grouse Influenced By Energy Development, Christopher P. Kirol, Andrew L. Sutphin, Laura Bond, Mark R. Fuller, Thomas L. Maechtle

Biology Faculty Publications and Presentations

Sagebrush Artemisia spp. habitats being developed for oil and gas reserves are inhabited by sagebrush obligate species--including the greater sage-grouse Centrocercus urophasianus (sage-grouse) that is currently being considered for protection under the U.S. Endangered Species Act. Numerous studies suggest increasing oil and gas development may exacerbate species extinction risks. Therefore, there is a great need for effective on-site mitigation to reduce impacts to co-occurring wildlife such as sage-grouse. Nesting success is a primary factor in avian productivity and declines in nesting success are also thought to be an important contributor to population declines in sage-grouse. From 2008 to 2011 we …

Isolation Of Diverse Members Of The Aquificales From Geothermal Springs In Tengchong, China, Brian P. Hedlund, Anna-Louise Reysenbach, Liuquin Huang, John C. Ong, Zizhang Liu, Jeremy A. Dodsworth, Reham Ahmed, Amanda J. Williams, Brandon R. Briggs, Yitai Liu, Weiguo Hou, Hailiang Dong

Biology Faculty Publications and Presentations

The order Aquificales (phylum Aquificae) consists of thermophilic and hyperthermophilic bacteria that are prominent in many geothermal systems, including those in Tengchong, Yunnan Province, China. However, Aquificales have not previously been isolated from Tengchong. We isolated five strains of Aquificales from diverse springs (temperature 45.2–83.3°C and pH 2.6–9.1) in the Rehai Geothermal Field from sites in which Aquificales were abundant. Phylogenetic analysis showed that four of the strains belong to the genera Hydrogenobacter, Hydrogenobaculum, andSulfurihydrogenibium, including strains distant enough to likely justify new species ofHydrogenobacter and Hydrogenobaculum. The additional strain may represent a …

Gene Expression Patterns That Support Novel Developmental Stress Buffering In Embryos Of The Annual Killifish Austrofundulus Limnaeus, Josiah T. Wagner, Jason E. Podrabsky

Biology Faculty Publications and Presentations

Background: The cellular signaling mechanisms and morphogenic movements involved in axis formation and gastrulation are well conserved between vertebrates. In nearly all described fish, gastrulation and the initial patterning of the embryonic axis occur concurrently with epiboly. However, annual killifish may be an exception to this norm. Annual killifish inhabit ephemeral ponds in South America and Africa and permanent populations persist by the production of stress-tolerant eggs. Early development of annual killifish is unique among vertebrates because their embryonic blastomeres disperse randomly across the yolk during epiboly and reaggregate several days later to form the embryo proper. In addition, annual …

Anthropogenic Noise Alters Bat Activity Levels And Echolocation Calls, Jessie P. Bunkley, Christopher J.W. Mcclure, Nathan J. Kleist, Clinton D. Francis, Jesse R. Barber

Biology Faculty Publications and Presentations

Negative impacts from anthropogenic noise are well documented for many wildlife taxa. Investigations of the effects of noise on bats however, have not been conducted outside of the laboratory. Bats that hunt arthropods rely on auditory information to forage. Part of this acoustic information can fall within the spectrum of anthropogenic noise, which can potentially interfere with signal reception and processing. Compressor stations associated with natural gas extraction produce broadband noise 24 hours a day, 365 days a year. With over half a million producing gas wells in the U.S. this infrastructure is a major source of noise pollution across …

Fearscapes: Mapping Functional Properties Of Cover For Prey With Terrestrial Lidar, Peter J. Olsoy, Jennifer S. Forbey, Janet L. Rachlow, Jordan D. Nobler, Nancy F. Glenn, Lisa A. Shipley

Biology Faculty Publications and Presentations

Heterogeneous vegetation structure can create a variable landscape of predation risk—a fearscape—that influences the use and selection of habitat by animals. Mapping the functional properties of vegetation that influence predation risk (e.g., concealment and visibility) across landscapes can be challenging. Traditional ground-based measures of predation risk are location specific and limited in spatial resolution. We demonstrate the benefits of terrestrial laser scanning (TLS) to map the properties of vegetation structure that shape fearscapes. We used TLS data to estimate the concealment of prey from multiple vantage points, representing predator sightlines, as well as the visibility of potential predators from the …

Effects Of Sex And Mycorrhizal Fungi On Gas Exchange In The Dioecious Salt Marsh Grass Distichlis Spicata, Kassandra Reuss-Schmidt, Todd N. Rosenstiel, Sally R. Rogers, Allie G. Simpson, Sarah M. Eppley

Biology Faculty Publications and Presentations

Premise of research. In dioecious plant species, males and females often differ in physiology, and mycorrhizal fungal relationships are likely to influence these differences. However, few data are available on the potential role of mycorrhizal fungi in altering sex-specific physiology and population sex ratios of dioecious plant species.

Methodology. In this study, we measured leaf gas exchange in a multifactorial greenhouse experiment with and without mycorrhizal fungal additions and under field conditions in Distichlis spicata, a dioecious C4 salt marsh grass, displaying extreme spatial sex ratio variation.

Pivotal results. We found a significant interaction between gas exchange, plant sex, and …

Extreme Tolerance And Developmental Buffering Of Uv-C Induced Dna Damage In Embryos Of The Annual Killifish Austrofundulus Limnaeus, Josiah Tad Wagner, Jason E. Podrabsky

Biology Faculty Publications and Presentations

Free-living aquatic embryos are often at risk of exposure to ultraviolet radiation (UV-R). Successful completion of embryonic development depends on efficient removal of DNA lesions, and thus many aquatic embryos have mechanisms to reverse DNA lesions induced by UV-R. However, little is known of how embryos that are able to enter embryonic dormancy may respond to UV-R exposure and subsequent DNA damage. Embryos of the annual killifish Austrofundulus limnaeus are unique among vertebrates because their normal embryonic development includes (1) a complete dispersion of embryonic blastomeres prior to formation of the definitive embryonic axis, and (2) entry into a state …

A Coxiella-Like Endosymbiont Is A Potential Vitamin Source For The Lone Star Tick, Todd A. Smith, Timothy Driscoll, Joseph J. Gillespie, Rahul Raghavan

Biology Faculty Publications and Presentations

Amblyomma americanum (Lone star tick) is an important disease vector in the United States. It transmits several human pathogens, including the agents of human monocytic ehrlichiosis, tularemia, and southern tick-associated rash illness. Blood-feeding insects (Class Insecta) depend on bacterial endosymbionts to provide vitamins and cofactors that are scarce in blood. It is unclear how this deficiency is compensated in ticks (Class Arachnida) that feed exclusively on mammalian blood. A bacterium related to Coxiella burnetii, the agent of human Q fever, has been observed previously within cells of A. americanum. Eliminating this bacterium (CLEAA, Coxiella-like endosymbiont of A. americanum) with antibiotics …

Antibacterial Gene Transfer Across The Tree Of Life, Jason A. Metcalf, Lisa J. Funkhouser-Jones, Kristen A. Brileya, Anna-Louise Reysenbach, Seth R. Bordenstein

Biology Faculty Publications and Presentations

Though horizontal gene transfer (HGT) is widespread, genes and taxa experience biased rates of transferability. Curiously, independent transmission of homologous DNA to archaea, bacteria, eukaryotes, and viruses is extremely rare and often defies ecological and functional explanations. Here, we demonstrate that a bacterial lysozyme family integrated independently in all domains of life across diverse environments, generating the only glycosyl hydrolase 25 muramidases in plants and archaea. During coculture of a hydrothermal vent archaeon with a bacterial competitor, muramidase transcription is upregulated. Moreover, recombinant lysozyme exhibits broad-spectrum antibacterial action in a dose-dependent manner. Similar to bacterial transfer of antibiotic resistance genes, …

Using Phylogenetically-Informed Annotation (Pia) To Search For Light-Interacting Genes In Transcriptomes From Non-Model Organisms, Daniel L. Speiser, Molly S. Pankey, Alexander K. Zaharoff, Barbara A. Battelle, Heather D. Bracken-Grissom, Jesse W. Breinholt, Seth M. Bybee, Thomas W. Cronin, Anders Garm, Annie R. Lindgren, Nipam H. Patel, Megan L. Porter, Meredith E. Protas, Ajna S. Rivera, Jeanne M. Serb, Kirk S. Zigler, Keith A. Crandall, Todd H. Oakley

Biology Faculty Publications and Presentations

Background: Tools for high throughput sequencing and de novo assembly make the analysis of transcriptomes (i.e. the suite of genes expressed in a tissue) feasible for almost any organism. Yet a challenge for biologists is that it can be difficult to assign identities to gene sequences, especially from non-model organisms. Phylogenetic analyses are one useful method for assigning identities to these sequences, but such methods tend to be time-consuming because of the need to re-calculate trees for every gene of interest and each time a new data set is analyzed. In response, we employed existing tools for phylogenetic analysis to …

Choroid Plexus In Developmental And Evolutionary Perspective, Brent Roy Bill, Vladimir Korzh

Biology Faculty Publications and Presentations

The blood-cerebrospinal fluid boundary is present at the level of epithelial cells of the choroid plexus. As one of the sources of the cerebrospinal fluid (CSF), the choroid plexus (CP) plays an important role during brain development and function. Its formation has been studied largely in mammalian species. Lately, progress in other model animals, in particular the zebrafish, has brought a deeper understanding of CP formation, due in part to the ability to observe CP development in vivo. At the same time, advances in comparative genomics began providing information, which opens a possibility to understand further the molecular mechanisms involved …

The Dilemma Of Foraging Herbivores: Dealing With Food And Fear, Clare Mcarthur, Peter B. Banks, Rudy Boonstra, Jennifer Sorensen Forbey

Biology Faculty Publications and Presentations

For foraging herbivores, both food quality and predation risk vary across the landscape. Animals should avoid low-quality food patches in favour of high-quality ones, and seek safe patches while avoiding risky ones. Herbivores often face the foraging dilemma, however, of choosing between high-quality food in risky places or low-quality food in safe places. Here, we explore how and why the interaction between food quality and predation risk affects foraging decisions of mammalian herbivores, focusing on browsers confronting plant toxins in a landscape of fear. We draw together themes of plant–herbivore and predator–prey interactions, and the roles of animal ecophysiology, behaviour …

Mercury And Selenium In Stranded Indo-Pacific Humpback Dolphins And Implications For Their Trophic, Duan Gui, Ri-Qing Yu, Laiguo Chen, Laiguo Chen, Qin Tu, Hui Mo, Yuping Wu

Biology Faculty Publications and Presentations

As top predators in the Pearl River Estuary (PRE) of China, Indo-Pacific humpback dolphins (Sousa chinensis) are bioindicators for examining regional trends of environmental contaminants in the PRE. We examined samples from stranded S. chinensis in the PRE, collected since 2004, to study the distribution and fate of total mercury (THg), methylmercury (MeHg) and selenium (Se) in the major tissues, in individuals at different ages and their prey fishes from the PRE. This study also investigated the potential protective effects of Se against the toxicities of accumulated THg. Dolphin livers contained the highest concentrations of THg (32.34658.98 mg g21 dw) …

Solenopsis Invicta Virus (Sinv-1) Infection And Insecticide Interactions In The Red Imported Fire Ant (Hymenoptera: Formicidae), Danielle M. Tufts, Wayne B. Hunter, Blake Bextine

Biology Faculty Publications and Presentations

The red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae), is of great concern because of its destructive nature to endemic wildlife, livestock, and people. Various methods for managing this pest are currently being developed, including the use of viruses as biological control agents. In this study, the effectiveness of the Solenopsis invicta virus (SINV-1), (a positive sense, single-stranded RNA virus in the Dicistroviridae family (Genus: Aparavirus) which only infects the Genus Solenopsis) as an effective biological control agent against S. invicta infestation combination with commonly used insecticides was investigated. Surprisingly, ants treated with the virus experienced significantly greater survival …

Redescription Of Atopospira Galeata (Kahl, 1927) Nov. Comb. And A. Violacea (Kahl, 1926) Nov. Comb. With Redefinition Of Atopospira Jankowski, 1964 Nov. Stat. And Brachonella Jankowski, 1964 (Ciliophora, Armophorida), William A. Bourland, Laura Wendell

Biology Faculty Publications and Presentations

The taxonomy of the Metopidae (Ciliophora, Armophorida) remains poorly understood since most of its members have not been studied by modern morphologic and molecular methods. Recent molecular investigations have indicated that the two most species-rich genera, Metopus and Brachonella, are likely nonmonophyletic with at least one well-supported 18S rDNA clade comprised of a species from each of these genera (Brachonella galeata and Metopus violaceus). We investigated these two species with silver impregnation and scanning electron microscopy. Both taxa share important morphologic characteristics not described in other species of Metopus or Brachonella. These synapomorphies include: (1) a …

An Observation Of Apparent Teaching Behavior In The Pallid Bat, Antrozous Pallidus, Jessie P. Bunkley, Jesse R. Barber

Biology Faculty Publications and Presentations

During a laboratory study of pallid bat (Antrozous pallidus) hunting behavior, we observed an interaction wherein an adult female appeared to aid a juvenile male in learning a novel foraging task. This single observation adheres to the 3 requirements of teaching outlined by Caro and Hauser (1992). A female bat experienced with a hunting task modified her behavior in the presence of a naïve observing male, resulting in a cost of reduced food availability to the female when she was hungry, while directing the male to food resources and accelerating his learning of a foraging task. …

Movements Of A Juvenile Crowned Eagle (Harpyhaliaetus Coronatus) Tracked By Satellite Telemetry In Central Argentina, Vicente Urios, Maria Pilar Donat-Torres, Marc J. Bechard, Miguel Ferrer

Biology Faculty Publications and Presentations

Background: A juvenile Crowned Eagle was tagged at its nest with a satellite transmitter. The Crowned Eagle (Harpyhaliaetus coronatus) is one of the most unknown raptor species from the American continent. Their current distribution ranges from central Brazil to central Argentina, with a total population of 350–1500 individuals across this large area, being thus largely fragmented.

Results: During the three years of tracking the bird concentrated its movements in a range spanning for 12845 km², but concentrating mainly in four smaller areas accounting for 3073 km². The locations were recorded mainly over shrubland habitats …