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Full-Text Articles in Behavioral Neurobiology

Prenatal Acoustic Programming Of The Developing Hypothalamus: A Multi-Omic Analysis Of Heat Call Exposure In Zebra Finch Embryos, Prakrit Subba Aug 2026

Prenatal Acoustic Programming Of The Developing Hypothalamus: A Multi-Omic Analysis Of Heat Call Exposure In Zebra Finch Embryos, Prakrit Subba

All Dissertations

Climate change is intensifying thermal stress on wildlife globally, yet the developmental mechanisms by which embryos use prenatal cues to adjust to predicted environments remain unclear. In Australian zebra finches (Taeniopygia guttata), incubating parents produce distinctive heat call vocalizations at high ambient temperatures, and embryos exposed to these calls during late incubation develop altered growth trajectories, thermoregulatory capacity, and reproductive outcomes, suggesting a form of anticipatory developmental programming triggered by sound alone. To investigate the molecular basis of this response, I initially hypothesized that prenatal heat call exposure would act primarily through hypothalamic neuroendocrine reprogramming, given the central …


Structure And Diversity Of Play Behavior In Juvenile Belding's Ground Squirrels, Paul O'Driscoll Ii May 2026

Structure And Diversity Of Play Behavior In Juvenile Belding's Ground Squirrels, Paul O'Driscoll Ii

Master's Theses

Play behavior is ubiquitous among juvenile mammals and can promote motor, social, and cognitive development, providing multiple adaptive benefits to young animals in some cases. The structure of play can be linked to the benefits it confers on individuals. I evaluated the structure of play in juvenile Belding’s ground squirrels (Urocitellus beldingi), a species for which multiple adaptive benefits of play have been identified. I compared juvenile play in U. beldingi to adult sexual and aggressive behavior to determine and classify similarities between juvenile play and adult behavior. I evaluated diversity in the behaviors expressed during play interactions and diversity …


Beat Behind The Bark: Investigating The Effects Of Music Tempo On Canine Physiological And Psychological States, Marley Lanier May 2026

Beat Behind The Bark: Investigating The Effects Of Music Tempo On Canine Physiological And Psychological States, Marley Lanier

Honors Theses

This study explores how the tempo of music, measured in beats per minute (BPM), affects dogs' heart rate and oxytocin levels, adding to past research that shows that music affects physiological and behavioral states in dogs. Being able to understand these effects holds practical implications for improving overall well-being in canines in environments such as homes, shelters, and clinical environments. Using a within-subjects design, the dogs were exposed to slow and fast versions of one instrumental track, along with a white noise baseline condition before each randomized condition. Heart rate was recorded continuously throughout the session, and oxytocin saliva samples …


Factors Influencing Sex-Based Movement Patterns In The Banded Garden Spider, Argiope Trifasciata (Forskal, 1776), Brandon R. Marrone Apr 2026

Factors Influencing Sex-Based Movement Patterns In The Banded Garden Spider, Argiope Trifasciata (Forskal, 1776), Brandon R. Marrone

School of Biological Sciences: Dissertations, Theses, and Student Research

Movement is a fundamental ecological process shaping how individuals acquire mates, secure resources, and navigate heterogeneous landscapes. In species with extreme sexual size dimorphism (eSSD), such as the orb‑weaving spider Argiope trifasciata, males and females are predicted to experience different movement pressures driven by their reproductive roles. This thesis examines how individual traits and environmental conditions jointly influence movement across both sexes. Across two field seasons, we used intensive mark–recapture methods to quantify natural movement patterns of males and females within mapped prairie plots. The first chapter tested long‑standing predictions gravity hypothesis, which posits that smaller males benefit from …


The Evolution Of Self-Awareness In Conscious Organisms, Robert A. Dielenberg Mar 2026

The Evolution Of Self-Awareness In Conscious Organisms, Robert A. Dielenberg

Journal of Conscious Evolution

Self-awareness in conscious organisms remains an enduring puzzle for philosophers and neuroscientists. This paper proposes a tripartite evolutionary model tracing its emergence through three sequential types: the sensory self (primitive egocentric, valence-tagged awareness of sensory states), the bodily self (external objectification of the body as a distinct entity, evidenced by mirror self-recognition in select species), and the cognitive self (internal objectification of the mind enabling recursive self-transformation). Adopting a monist framework, the model treats consciousness as the biological substrate of delayed neural apperception (the “Delayed Present”), within which selves emerge as increasingly sophisticated models. It distinguishes non-human metacognition—implicit, online regulation—from …


Adapting To Uncertainty: Effects Of Environmental Instability On Zebrafish Behaviour, Britney Sekulovski Jan 2026

Adapting To Uncertainty: Effects Of Environmental Instability On Zebrafish Behaviour, Britney Sekulovski

Theses and Dissertations (Comprehensive)

This thesis addresses how animals cope with unpredictability by testing how environmental stability changes the behaviour of zebrafish (Danio rerio) in two complementary experiments. In Experiment 1, fish lived for three months in either a Dynamic environment (water temperature, feeding times, habitat complexity, group size, and membership fluctuated) or a Stable environment (all factors constant). Across seven behavioural assays, measuring individual behavioural differences, collective movement, social memory, and information use, Dynamic condition fish were found to be more sensitive to social cues and more likely to conform to recent social information, independent of the quality of the information …


Investigating The Impact Of Psychoactive Plant Alkaloids On Bumblebee Behavior, Skylar Mathieson Aug 2025

Investigating The Impact Of Psychoactive Plant Alkaloids On Bumblebee Behavior, Skylar Mathieson

Electronic Theses and Dissertations

The relationship between plants and their pollinators is often framed as a food-for-service exchange in which plants provide pollinators with nutrients in exchange for transport of their gametes. However, in addition to essential nutrients, many floral rewards (nectar, pollen) also contain defensive toxins such as alkaloids, with largely unexplored consequences for plant-pollinator interactions. I investigated how three psychoactive and addictive plant alkaloids — caffeine, nicotine, and morphine — affect foraging behavior in generalist bumblebees (Bombus impatiens) using robotic flowers. Each of these chemicals affected pollinator foraging behavior, influencing the number and duration of their visits to flowers in …


Investigating Prefrontal Cortex Neuronal Signatures Of Opioid-Induced Risk-Taking Behavior, Cana Quave May 2025

Investigating Prefrontal Cortex Neuronal Signatures Of Opioid-Induced Risk-Taking Behavior, Cana Quave

Dissertations and Theses (Open Access)

Opioid use disorder occurs alongside impaired risk-related decision-making, but the underlying neural correlates are unclear. We developed an approach-avoidance conflict task using a modified conditioned place preference procedure to study neural signals of risky opioid seeking in the prefrontal cortex, a region implicated in executive decision-making. Following morphine conditioned place preference, rats underwent a conflict test in which fear-inducing cat odor was introduced in the previously drug-paired side of the apparatus. While the saline-exposed control group avoided cat odor, the morphine group included two subsets of rats that either maintained a preference for the paired side despite the presence of …


Effects Of Retinal Gonadotropin-Releasing Hormone On Reproductive Behavior And Retinal Activity In The Cichlid Fish, Astatotilapia Burtoni, Chase Anselmo Oct 2024

Effects Of Retinal Gonadotropin-Releasing Hormone On Reproductive Behavior And Retinal Activity In The Cichlid Fish, Astatotilapia Burtoni, Chase Anselmo

LSU Master's Theses

Many animals use visual signals to evade predators, forage for food, and communicate with conspecifics. Some animals that shift into and out of breeding stages, such as fish, can modulate their visual sensitivity to better adapt to changing internal and external conditions. Previous work in our lab demonstrated that females of the cichlid fish Astatotilapia burtoni have improved visual sensitivity when they enter reproductive readiness. However, little is known about how fish and vertebrates more broadly modulate visual sensitivity across their reproductive cycle. We hypothesized that the hormone and neuromodulator gonadotropin-releasing hormone (GnRH) could modulate visual sensitivity because GnRH is …


Compensating For Effects Of Autotomy On Locomotion In Stink Bugs (Halyomorpha Halys), Lilian C. Jubb Oct 2023

Compensating For Effects Of Autotomy On Locomotion In Stink Bugs (Halyomorpha Halys), Lilian C. Jubb

Longwood Senior Thesis Proposal

Autotomy is the voluntary release of an appendage, typically to avoid predation or entrapment. Autonomy of a single leg can have minimal effects on locomotion in arthropods with more than six legs. Hexapods, however, use a double-tripod gait which is completely disrupted with the loss of a single leg, thus locomotor performance should be greatly reduced leading to lower survival. Autotomy is so costly to locomotion that some insects regenerate a lost limb in a subsequent molt leading to reduced investment in other important functions (e.g. reproduction). Autotomy persists in adult hexapods with no opportunity for regeneration, which suggests some …


Statistical And Biological Analyses Of Acoustic Signals In Estrildid Finches, Moises Rivera Jun 2023

Statistical And Biological Analyses Of Acoustic Signals In Estrildid Finches, Moises Rivera

Dissertations, Theses, and Capstone Projects

Acoustic communication is a process that involves auditory perception and signal processing. Discrimination and recognition further require cognitive processes and supporting mechanisms in order to successfully identify and appropriately respond to signal senders. Although acoustic communication is common across birds, classical research has largely disregarded the perceptual abilities of perinatal altricial taxa. Chapter 1 reviews the literature of perinatal acoustic stimulation in birds, highlighting the disproportionate focus on precocial birds (e.g., chickens, ducks, quails). The long-held belief that altricial birds were incapable of acoustic perception in ovo was only recently overturned, as researchers began to find behavioral and physiological evidence …


Modulatory Effects Of Schreckstoff On The Startle Escape Response Of Goldfish (Carassius Auratus), Denis Shor Jun 2023

Modulatory Effects Of Schreckstoff On The Startle Escape Response Of Goldfish (Carassius Auratus), Denis Shor

Dissertations, Theses, and Capstone Projects

Fish face a variety of stressors in nature, a prominent one being a predatory threat to which they respond with an adaptive behavioral response thus maximizing their chances of survival. A primary anti-predatory behavior is a startle response or C-start. This behavior is controlled by two large reticulospinal neurons, the Mauthner cells (M-cells) which integrate multimodal sensory inputs. Detecting a predator involves various sensory information such as visual and mechanosensory cues but also olfaction. Indeed, following an attack, an injured fish will release an alarm pheromone (Schreckstoff) and thus acutely alert close by conspecifics. However, whether the alarm pheromone in …


Nicotiana Glauca Graham (Solanaceae) Bioactivity And Toxic Effects On Mortality, Feeding Behavior And Pupation Choice Of Drosophila Melanogaster Larvae (Diptera: Drosophilidae), Abir Bouzar, Sarra Habbachi, Ibtissem Samai, Makarem Rahat, Nour El Imene Boublata, Saadane Fatma Zohra, Wafa Habbachi, Benhissen Saliha, Abdelkrim Tahraoui Dec 2022

Nicotiana Glauca Graham (Solanaceae) Bioactivity And Toxic Effects On Mortality, Feeding Behavior And Pupation Choice Of Drosophila Melanogaster Larvae (Diptera: Drosophilidae), Abir Bouzar, Sarra Habbachi, Ibtissem Samai, Makarem Rahat, Nour El Imene Boublata, Saadane Fatma Zohra, Wafa Habbachi, Benhissen Saliha, Abdelkrim Tahraoui

Journal of Bioresource Management

Nicotiana glauca is a medicinal plant used by traditional healers as antibacterial, antifungal, antiviral and anti-inflammatory medicines. The leaves of N. glauca are very rich in indolic alkaloids which give it a larvicidal power which allows us to use it as a bio-insecticide.In the present study we were perfomed in the direct (mortality) and indirect (food attractiveness and pupation) toxic effects of the aqueous extract of N. glauca on the mortality and feeding behavior of the fruit fly Drosophila melanogaster. The treatment was administered to 2nd instar larvae (L2) and the mortality rate was monitored for 15 days. Similarly we …


Use Of Intramodal Odor Cues In Shelter Recognition By An Amblypygid, Nathan Bostelman Apr 2022

Use Of Intramodal Odor Cues In Shelter Recognition By An Amblypygid, Nathan Bostelman

Honors Projects

Amblypygids are nocturnal arthropods which live in cluttered habitats and possess the ability to navigate home after displacements of up to ten meters. Homing amblypygids rely on olfactory and tactile information gathered by their antenniform legs to navigate successfully. Given that odor signals encountered in nature are complex and dynamic, navigation via olfaction presents unique challenges related to signal uncertainty. To understand how amblypygids perceive perceive complex odors and what they learn from them, individuals of the subtropical amblypygid P. marginemaculatus were trained to associate a blend of two odors with a shelter and tested on three treatments: the reinforced …


Sentience In Decapods: Difficulties To Surmount, Michael L. Woodruff Jan 2022

Sentience In Decapods: Difficulties To Surmount, Michael L. Woodruff

Animal Sentience

In the target article Crump et al. present 8 criteria to assess whether decapods experience pain. Four of these -- sensory integration, motivational trade-offs, flexible self-protection, and associative learning -- could be used to assess sentience in general. In this commentary I discuss difficulties with using these criteria to provide evidence of sentience in decapods, particularly if this evidence is to change public opinion and policies. These difficulties are lack of evidence, the potential to eventually explain the neurobiological basis of the behaviors chosen as criteria, thereby eliminating any explanatory work for sentience, and the reluctance to bring animals that …


A Framework For Evaluating Evidence Of Pain In Animals, Matilda Gibbons, Lars Chittka Jan 2022

A Framework For Evaluating Evidence Of Pain In Animals, Matilda Gibbons, Lars Chittka

Animal Sentience

Crump et al. define eight criteria indicating sentience in animals, with a focus on pain. Here, we point out the risk of false negative or false positive diagnoses of pain. Criteria of different levels of inclusivity are useful for using the precautionary principle in animal welfare considerations, and for more formal scientific evidence of pain. We suggest tightening the criteria -- from more general evidence of sentience to pain alone -- because crucial evidence for animal welfare decisions might otherwise be missed for animals subjected to invasive and injurious procedures.


Neurotranscriptomic Changes Associated With Chick-Directed Parental Care In Adult Non-Reproductive Japanese Quail, Patricia C. Lopes, Robert De Brujin Jul 2021

Neurotranscriptomic Changes Associated With Chick-Directed Parental Care In Adult Non-Reproductive Japanese Quail, Patricia C. Lopes, Robert De Brujin

Biology, Chemistry, and Environmental Sciences Faculty Articles and Research

For many species, parental care critically affects offspring survival. But what drives animals to display parental behaviours towards young? In mammals, pregnancy-induced physiological transformations seem key in preparing the neural circuits that lead towards attraction (and reduced-aggression) to young. Beyond mammalian maternal behaviour, knowledge of the neural mechanisms that underlie young-directed parental care is severely lacking. We took advantage of a domesticated bird species, the Japanese quail, for which parental behaviour towards chicks can be induced in virgin non-reproductive adults through a sensitization procedure, a process that is not effective in all animals. We used the variation in parental responses …


Effects Of Non-Photic Zeitgebers On The Circadian Clock In The Common House Spider, Parasteatoda Tepidariorum (Araneae: Theridiidae), Mattea Garmany, Darrell Moore, Thomas C. Jones May 2020

Effects Of Non-Photic Zeitgebers On The Circadian Clock In The Common House Spider, Parasteatoda Tepidariorum (Araneae: Theridiidae), Mattea Garmany, Darrell Moore, Thomas C. Jones

Undergraduate Honors Theses

Circadian rhythms are endogenous cycles that control physiological and behavioral changes that can be affected by environmental factors which allow most eukaryotic organisms to synchronize their daily activities with the 24-hour day. Parasteatoda tepidariorum,the common house spider, demonstrates a short-period circadian clock averaging 21.6 hours when left in constant darkness, yet they are able to entrain to a 24-hour light cycle. We tested whether these spiders were able to use non-photic Zeitgebers to entrain to the 24-hour day. Periodic presentation of food and disturbance were not found to be effective cues for the spiders’ entrainment. A few individuals were clearly …


Behavioral Ecology Of A Desert Ambush Predator: Assessing Movement Patterns, Habitat And Microhabitat Use, And The Innate Feeding Response Of Eastern Black-Tailed Rattlesnakes (Crotalus Ornatus) In The Northern Chihuahuan Desert, James Emerson Jan 2020

Behavioral Ecology Of A Desert Ambush Predator: Assessing Movement Patterns, Habitat And Microhabitat Use, And The Innate Feeding Response Of Eastern Black-Tailed Rattlesnakes (Crotalus Ornatus) In The Northern Chihuahuan Desert, James Emerson

Open Access Theses & Dissertations

Research on the ecology and evolution of rattlesnakes has been sporadic over the past 80 years, but has seen recent expansion into the diverse fields of physiology, physiological ecology, life history, behavioral ecology, ethology, reproductive biology, chemical ecology, venom biochemistry and medicine, conservation, and many other subdisciplines. The development of small, implantable VHF radiotransmitters in the 1980s revolutionized research in the field of behavioral ecology for rattlesnakes, which are uniquely suited for radiotelemetry studies because they possess several morphological, physiological, and behavioral characteristics that are unique among terrestrial vertebrates. The widespread application of radiotelemetry by the mid-1980s advanced the model …


Tribal Brains In The Global Village: Deeper Roots Of The Pandemic, Robert Gerlai Jan 2020

Tribal Brains In The Global Village: Deeper Roots Of The Pandemic, Robert Gerlai

Animal Sentience

I briefly recap the messages of the target article by Wiebers & Feigin (2020) and the accompanying peer commentaries about what we learn from the COVID-19 pandemic. Using the rapid evolution of viruses as an example of the importance of prevention, I explore why it is difficult for our species to foresee and prevent unintended global changes resulting from human activity. I end with a discussion about the long-term future, the ultimate problem inherent in our current mindset and the structure of our economy: growth.


Inhibition Of Pain Or Response To Injury In Invertebrates And Vertebrates, Matilda Gibbons, Sajedeh Sarlak Jan 2020

Inhibition Of Pain Or Response To Injury In Invertebrates And Vertebrates, Matilda Gibbons, Sajedeh Sarlak

Animal Sentience

In certain situations, insects appear to lack a response to noxious stimuli that would cause pain in humans. For example, from the fact that male mantids continue to mate while being eaten by their partner it does not follow that insects do not feel pain; it could be the result of modulation of nociceptive inputs or behavioural outputs. When we try to infer the underlying mental state of an insect from its behaviour, it is important to consider the behavioural effects of the associated physiological and neurobiological mechanisms.


The ‘Thing’ From This World, Sergio M. Pellis Jan 2019

The ‘Thing’ From This World, Sergio M. Pellis

Animal Sentience

Science progresses by making contrasts, and the living world is a gold mine of contrasts. Often disciplines become victims by focusing on too narrow a slice of that diversity, leading to a myopic view of how nature works. The relationships between the brain and behavior have been intensively studied in vertebrates, especially mammals, and we have become complacent in our assumptions about how behavior is constructed. As the target article by Mather (2019) shows, the relationship between the brain and behavior in octopuses forces us to reevaluate some of those assumptions.


Behavioral Insights Into Nociceptor Function: A Systematic Approach To Understanding Postsurgical And Neuropathic Pain Mechanisms In Rats, Max Odem Dec 2018

Behavioral Insights Into Nociceptor Function: A Systematic Approach To Understanding Postsurgical And Neuropathic Pain Mechanisms In Rats, Max Odem

Dissertations and Theses (Open Access)

Postsurgical and neuropathic pain are each clinically common, and often associated with ongoing pain. Ongoing pain has been linked to ongoing activity (OA) in human C-fiber nociceptors. Preclinical studies using rodent neuropathic models have concentrated on allodynia driven by OA generated in non-nociceptive Aβ fibers, but little attention has been paid to postsurgical pain in sham controls or to C-fiber nociceptor OA promoting ongoing pain.

Operant assays that reveal negative motivational and cognitive aspects of voluntary pain-related behavior may be particularly sensitive to pain-related alterations. In the mechanical conflict (MC) test, rodents can freely choose to escape from a brightly …


Diel Rhythmicity Found In Behavior But Not Biogenic Amine Levels In The Funnel-Web Spider Agelenopsis Pennsylvanica (Araneae, Agelenidae), Alexander E. Demarco May 2018

Diel Rhythmicity Found In Behavior But Not Biogenic Amine Levels In The Funnel-Web Spider Agelenopsis Pennsylvanica (Araneae, Agelenidae), Alexander E. Demarco

Electronic Theses and Dissertations

Quantifying individual differences in behavior and the extent that behavior is influenced by circadian control is of paramount importance in behavioral ecology. In addition, the proximate mechanisms underlying behavior are also critical in order to obtain a more complete picture of how behavior evolves. Biogenic amines (BAs) are simple nitrogenous compounds derived from amino acids and have been consistently and extensively linked to behavior. For this study, we analyzed temporal patterns of BAs in relation to the antipredator (boldness) and aggressive behavior in female Agelenopsis pennsylvanica, a funnel-web spider. Using HPLC-ED, we compared behavioral responses to temporal patterns of …


Social Influences On Songbird Behavior: From Song Learning To Motion Coordination, Iva Ljubičić Feb 2018

Social Influences On Songbird Behavior: From Song Learning To Motion Coordination, Iva Ljubičić

Dissertations, Theses, and Capstone Projects

Social animals learn during development how to integrate successfully into their group. How do social interactions combine to maintain group cohesion? We first review how social environments can influence the development of vocal learners, such as songbirds and humans (Chapter 1). To bypass the complexity of natural social interactions and gain experimental control, we developed Virtual Social Environments, surrounding the bird with videos of manipulated playbacks. This way we were able to design sensory and social scenarios and test how social zebra finches adjust their behavior (Chapters 2 & 3). A serious challenge is that the color output of a …


Only The Human Brain Has The Cognitive Capacity For Jealousy, Donatella Marazziti Jan 2018

Only The Human Brain Has The Cognitive Capacity For Jealousy, Donatella Marazziti

Animal Sentience

Jealousy is exclusively a human phenomenon because nonhuman animals lack the brain structures regulating the higher processes underlying jealousy.


Sentience In Fishes: More On The Evidence, Michael L. Woodruff Jan 2018

Sentience In Fishes: More On The Evidence, Michael L. Woodruff

Animal Sentience

In my target article, I argued that the brains of ray-finned fishes of the teleost subclass (Actinopterygii) are sufficiently complex to support sentience — that these fishes have subjective awareness of interoceptive and exteroceptive sense experience. Extending previous theories centered on the tectum, I focused on the organization of the fish pallium. In this Response to the commentaries, I clarify that I do not propose that the fish pallium is, or must be, homologous to the mammalian neocortex to play a role in sentience. Some form of a functionalist approach to explaining the neural basis of sentience across taxa is …


Comparing Consistency Of Stress And Anxiety-Related Behaviors Across Time In Zebrafish (Danio Rerio), Matthew R. Baker, Alex Goodman Mar 2017

Comparing Consistency Of Stress And Anxiety-Related Behaviors Across Time In Zebrafish (Danio Rerio), Matthew R. Baker, Alex Goodman

UNO Student Research and Creative Activity Fair

Animals are frequently faced with stressors in their environment that they must overcome to survive and reproduce. Across vertebrates, two distinct stress coping styles or ‘personalities’ have been observed known as proactive (bold) and reactive (shy). Animal personalities may be advantageous by limiting individual variation and balancing different trade-offs in unpredictable environments. When identifying animal personalities, behavioral phenotypes must be consistent and repeatable across contexts and time. Here we use selectively bred lines of shy and bold zebrafish, previously shown to have consistent divergent fear- and anxiety-related behaviors across contexts, to test the repeatability and consistency of these behaviors across …


The Evolutionary History Of Consciousness, Eirik Søvik, Clint Perry Sep 2016

The Evolutionary History Of Consciousness, Eirik Søvik, Clint Perry

Animal Sentience

Klein & Barron argue that insects are capable of subjective experience, i.e., sentience. Whereas we mostly agree with the conclusion of their arguments, we think there is an even more important message to be learned from their work. The line of reasoning opened by Klein & Barron proves instructive for how neuroscientists can and should explore the biological phenomenon of consciousness.


Exploring The Relationship Between Behaviour And Neurochemistry In The Polyphenic Spider, Anelosimus Studiosus (Araneae: Theridiidae), Jennifer B. Price Aug 2016

Exploring The Relationship Between Behaviour And Neurochemistry In The Polyphenic Spider, Anelosimus Studiosus (Araneae: Theridiidae), Jennifer B. Price

Electronic Theses and Dissertations

The importance of social behaviour is evident in human society, but there are both costs and benefits associated with cooperation and sociality throughout the animal kingdom. At what point do the benefits outweigh the costs, and when do selective pressures favour sociality and colonization over solitude and independence? To investigate these questions, we have focused on an anomalous species of spider, Anelosimus studiosus, also known now as the northern social spider. Throughout its broad range, A. studiosus is solitary and aggressive, but recently, colonies of cooperative and social individuals have been observed at northern latitudes. This leads to two …