Cicada nymphs dominate American black bear diet in a desert riparian area.

American black bears are considered dependent on high-elevation forests or other montane habitats in the drylands of western North America. Black bear sign, including that of cubs, was observed throughout the summers of 2015, 2016, and 2018 along a perennial desert river in the Sonoran Desert of Arizona. We analyzed the contents of 21 black bear scats, collected from May to October of 2016 and 2018. Apache cicada nymphs (Diceroprocta apache) were the dominant food item, occurring in 90% of scats and comprising an average of 59% of scat contents. In the process of excavating these nymphs, bears created large areas of turned-over soil, a form of ecosystem engineering with potential implications for soils, vegetation, and fluvial geomorphology. Given that species distributions are shaped by physiological and ecological contexts, as well as anthropogenic legacies, it is possible that black bears once occurred more commonly in desert riparian systems prior to widespread agricultural development, hunting, and dewatering. Although more research is necessary, we suggest that desert riparian systems may be an alternative habitat for black bears. Better understanding the diet and habitat breadth of American black bears is important in the context of increasing landscape fragmentation and militarization in the U.S.-Mexican borderlands.

Destruction of Spiderwebs and Rescue of Ensnared Nestmates by a Granivorous Desert Ant (Veromessor pergandei).

Prey species rarely seek out and dismantle traps constructed by their predators. In the current study, we report an instance of targeted trap destruction by an invertebrate and a novel context for rescue behavior. We found that foragers of the granivorous desert ant (Veromessor pergandei) identify and cooperatively dismantle spiderwebs (Araneae: Theridiidae, Steatoda spp., and Asagena sp.) During group foraging, workers ensnared in webs are recovered by sisters, which transport them to the nest and groom away their silk bindings. The presence of an ensnared nestmate and chemical alarm signal significantly increased the probability of web removal and nestmate retrieval. A subset of larger-bodied foragers participated in web removal, and 6.3% became tangled or were captured by spiders. Most animals that perform rescue behavior live in small groups, but V. pergandei colonies include tens of thousands of short-lived workers. To maintain their size, large colonies must collect enough seeds to produce 650 new ants each day. We hypothesize that the removal of spiderwebs allows for an unimpeded income of seeds on a single foraging path during a brief daily temperature window. Despite the cost to individuals, webs are recognized and removed only when workers are captured in them.

Behavior and exocrine glands in the myrmecophilous beetle Dinarda dentata (Gravenhorst, 1806) (Coleoptera: Staphylinidae: Aleocharinae).

The nests of advanced eusocial ant species can be considered ecological islands with a diversity of ecological niches inhabited by not only the ants and their brood, but also a multitude of other organisms adapted to particular niches. In the current paper, we describe the myrmecophilous behavior and the exocrine glands that enable the staphylinid beetle Dinarda dentata to live closely with its host ants Formica sanguinea. We confirm previous anecdotal descriptions of the beetle's ability to snatch regurgitated food from ants that arrive with a full crop in the peripheral nest chambers, and describe how the beetle is able to appease its host ants and dull initial aggression in the ants.

The non-additive effects of body size on nest architecture in a polymorphic ant.

Like traditional organisms, eusocial insect societies express traits that are the target of natural selection. Variation at the colony level emerges from the combined attributes of thousands of workers and may yield characteristics not predicted from individual phenotypes. By manipulating the ratios of worker types, the basis of complex, colony-level traits can be reduced to the additive and non-additive interactions of their component parts. In this study, we investigated the independent and synergistic effects of body size on nest architecture in a seasonally polymorphic harvester ant, Veromessor pergandei Using network analysis, we compared wax casts of nests, and found that mixed-size groups built longer nests, excavated more sand and produced greater architectural complexity than single-sized worker groups. The nests built by polymorphic groups were not only larger in absolute terms, but larger than expected based on the combined contributions of both size classes in isolation. In effect, the interactions of different worker types yielded a colony-level trait that was not predicted from the sum of its parts. In nature, V. pergandei colonies with fewer fathers produce smaller workers each summer, and produce more workers annually. Because body size is linked to multiple colony-level traits, our findings demonstrate how selection acting on one characteristic, like mating frequency, could also shape unrelated characteristics, like nest architecture.This article is part of the theme issue 'Interdisciplinary approaches for uncovering the impacts of architecture on collective behaviour'.

Behavior and exocrine glands in the myrmecophilous beetle Lomechusoides strumosus (Fabricius, 1775) (formerly called Lomechusa strumosa) (Coleoptera: Staphylinidae: Aleocharinae).

To become integrated into an ant society, myrmecophilous parasites must overcome both the defenses and the communication system of their hosts. Some aleocharine staphylinid beetles employ chemical and tactile strategies to invade colonies, where they later consume ant brood and participate in parasitic trophallaxis with host ants. By producing compounds that both appease their hosts and stimulate adoption, the beetles are able to live in and deposit their own eggs in the well defended ant nest. In the current paper, previous findings on the myrmecophilous behavior and morphological features of the staphylinid beetle Lomechusoides (formerly Lomechusa) strumosus are reviewed and re-evaluated. Hitherto unpublished results concerning the beetles' ability to participate in the social food flow of their host ants are reported. Furthermore, we present an analysis and documentation of the behavioral interactions between beetles and host ants during the adoption process, and we report new histological and scanning electron microscopic analyses of the exocrine glands and morphological adaptations that underlie the myrmecophilous behavior of L. strumosus. The main features of L. strumosus are compared with those of the staphilinid myrmecophile Lomechusa (formerly Atemeles) pubicollis. The paper concludes with a description of the life trajectory of L. strumosus and presents a brief history and discussion of the hypotheses concerning the evolution of myrmecophily in L. strumosus and other highly adapted myrmecophilous parasites.

Amphotis marginata (Coleoptera: Nitidulidae) a highwayman of the ant Lasius fuliginosus.

The space occupied by evolutionarily advanced ant societies can be subdivided into functional sites, such as broodchambers; peripheral nest chambers; kitchen middens; and foraging routes. Many predators and social parasites are specially adapted to make their living inside specific niches created by ants. In particular, the foraging paths of certain ant species are frequented by predatory and kleptoparasitic arthropods, including one striking example, the nitidulid beetle, Amphotis marginata. Adults of this species obtain the majority of their nutrition by acting as a kind of "highwayman" on the foraging trails of the ant Lasius fuliginosus, where they solicit regurgitation from food laden ant-workers by mimicking the ant's food-begging signals. Employing food labeled with the radio isotope 32P, we assessed the quantities of food the beetles siphoned-off of food-laden ants, and we investigated the site preferences, behavioral mechanisms and possible morphological adaptations underlying the food kleptoparasitism of A. marginata.

The Florida Harvester Ant, Pogonomyrmex badius, Relies on Germination to Consume Large Seeds.

The Florida harvester ant, Pogonomyrmex badius, is one of many ant species and genera that stores large numbers of seeds in damp, underground chambers for later consumption. A comparison of the sizes of seeds recovered from storage chambers with those of seed husks discarded following consumption revealed that the used seeds are far smaller than stored seeds. This difference in use-rate was confirmed in field and laboratory colonies by offering marked seeds of various sizes and monitoring the appearance of size-specific chaff. Because foragers collect a range of seed sizes but only open small seeds, large seeds accumulate, forming 70% or more of the weight of seed stores. Major workers increase the rates at which small and medium seeds are opened, but do not increase the size range of opened seeds. Experiments limiting ant access to portions of natural seed chambers showed that seeds germinate during storage, but that the ants rapidly remove them. When offered alongside non germinating seeds, germinating seeds were preferentially fed to larvae. The rate of germination during the annual cycle was determined by both burial in artificial chambers at various depths and under four laboratory temperatures. The germination rate depends upon the species of seed, the soil/laboratory temperature and/or the elapsed time. The seasonal soil temperature cycle generated germination patterns that vary with the mix of locally-available seeds. Taken together, exploitation of germination greatly increases the resources available to the ants in space and time. While the largest seeds may have the nutritional value of 15 small seeds, the inability of workers to open large seeds at will precludes them from rapid use during catastrophic events. The harvester ant's approach to seed harvesting is therefore two-pronged, with both immediate and delayed payoffs arising from the tendency to forage for a wide variety of seeds sizes.

Sequential Subterranean Transport of Excavated Sand and Foraged Seeds in Nests of the Harvester Ant, Pogonomyrmex badius.

During their approximately annual nest relocations, Florida harvester ants (Pogonomyrmex badius) excavate large and architecturally-distinct subterranean nests. Aspects of this process were studied by planting a harvester ant colony in the field in a soil column composed of layers of 12 different colors of sand. Quantifying the colors of excavated sand dumped on the surface by the ants revealed the progress of nest deepening to 2 m and enlargement to 8 L in volume. Most of the excavation was completed within about 2 weeks, but the nest was doubled in volume after a winter lull. After 7 months, we excavated the nest and mapped its structure, revealing colored sand deposited in non-host colored layers, especially in the upper 30 to 40 cm of the nest. In all, about 2.5% of the excavated sediment was deposited below ground, a fact of importance to sediment dating by optically-stimulated luminescence (OSL). Upward transport of excavated sand is carried out in stages, probably by different groups of ants, through deposition, re-transport, incorporation into the nest walls and floors and remobilization from these. This results in considerable mixing of sand from different depths, as indicated in the multiple sand colors even within single sand pellets brought to the surface. Just as sand is transported upward by stages, incoming seeds are transported downward to seed chambers. Foragers collect seeds and deposit them only in the topmost nest chambers from which a separate group of workers rapidly transports them downward in increments detectable as a "wave" of seeds that eventually ends in the seed chambers, 20 to 80 cm below the surface. The upward and downward transport is an example of task-partitioning in a series-parallel organization of work carried out by a highly redundant work force in which each worker usually completes only part of a multi-step process.

The roots of multilevel selection: concepts of biological individuality in the early twentieth century.

As multilevel selection theory has gained greater acceptance over the past quarter-century, scientists and scholars have shown an increased interest in the theory's historical antecedents. Despite this interest, however, the early twentieth century remains largely unexplored. It is generally assumed that biologists thought "naively" about evolutionary dynamics during this era, and that their attempts to explain biological phenomena often lacked sophistication. Now that several recent works have called attention to the complex relationship between biological individuality and the levels of selection, we believe it will prove instructive to revisit these early-twentieth-century biologists and reassess their criteria for biological individuality. Doing so reveals that they constructed a multilevel explanation of evolution that anticipated modern interpretations in several important ways. Though it is certainly true that most of these early biologists failed to recognize natural selection's pervasive agency, it is no less true that one of them, termite expert Alfred Emerson, artfully united the multilevel theory of "emergent evolution" with natural selection in a way that differs but little from the theory of multilevel selection that many scientists and scholars now promote. After reviewing the historical record, we place these early-twentieth-century biologists in their proper historical context, and we compare their interpretation of evolution with modern interpretations.