Soil texture as an influence on the distribution of the desert seed-harvester ants Pogonomyrmex rugosus and Messor pergandei

Summary Pogonomyrmex rugosus and Messor pergandei are ecologically similar species of desert seed-harvester ants that coexist in numerous areas throughout the Sonoran and Mohave Deserts. However, these two species also commonly segregate along physical gradients, with each species predominating in areas that differ in soil texture and/or topographic relief. Along gradients that included bajada and alluvial flat habitats, P. rugosus occurred alone in coarse-textured soils near mountains, while M. pergandei occurred alone in finer-textured soils further away. Conversely, along a vegetation gradient that included creosote bush and saltbush habitats, P. rugosus occurred alone in finer-textured soils than those occupied by either M. pergandei alone or both species in coexistence. However, in both situations clay content was significantly higher in areas occupied by P. rugosus alone, and at the latter site clay content was correlated with relative abundance of each species. Moreover, local distribution pattern of these two species may be related to the effects of clay on water retention, with retention being highest in areas occupied by P. rugosus alone. Differences in reproductive ecology may also affect these patterns as P. rugosus reproductive flights follow summer monsoon rains, while those of M. pergandei occur during the milder winter and spring.     

Water loss in desert ants: caste variation and the effect of cuticle abrasion

Water loss rates (WLRs) varied across castes (workers > alate males > alate females) for desert ants in the genera Aphaenogaster, Messor and Pogonomyrmex. Exposure to soil caused increased WLRs in workers and foundresses (mated, dealate females), apparently because of cuticular abrasion caused by nest excavation. Moreover, field‐collected workers and foundresses from incipient nests (those exposed to soil) had similar WLRs, as did unabraded workers and alate females (those unexposed to soil). These data call into question previous adaptive scenarios for differences in WLRs across ant species and castes. For live alate females, WLRs increased over two stages. The first increase occurred immediately after mating, and the second occurred for foundresses collected 2 days later from incipient nests. By contrast, WLRs of HCN‐killed females were unaffected by mating, but increased significantly for foundresses collected from 2‐day‐old nests.     

Differential learning and memory by co-occurring ant species

Foragers of the antsMessor pergandei andPogonomyrmex rugosus experience differing levels of variability in the distribution of seeds they harvest due to species-specific differences in foraging behavior.Messor pergandei foragers experience more variable seed distributions and densities, learn to recognize a novel seed faster but forget this information faster thanP. rugosus, which experiences more constant seed distributions even in the same habitat. Rate of learning to recognize a novel seed species was negatively associated with measures of seed species diversity for both ants.Messor pergandei foragers respond to variation in seed density by varying number of seeds handled per seed harvested, whileP. rugosus foragers do not. Memory of a novel seed exceeds forager longevity, due perhaps to use of seed caches as a type of information center.     

Complex foraging ecology of the red harvester ant and its effect on the soil seed bank

Granivory is an important interaction in the arid and semi-arid zones of the world, since seeds form an abundant and nutritious resource in these areas. While species of the genus Pogonomyrmex have been studied in detail as seed predators, their impact on seed abundance in the soil has not yet been explored in sufficient depth. We studied the impact of the harvesting activities of the ant Pogonomyrmex barbatus on seed abundance in the soil of the Zapotitlán valley, Mexico. We found that P. barbatus activity significantly impacts the abundance of seeds in the soil, which is lower in the sites where P. barbatus forages than it is in sites with no recorded foraging. We also found that P. barbatus distributes intact seeds of three tree species, two of which are nurse plants, and could consequently be promoting the establishment of these species. Using tools derived from graph theory, we observed that the ant-seed interactions exhibit a nested pattern; where more depredated seed species seem to be the more spatially abundant in the environment. This study illustrates the complex foraging ecology of the harvester ant P. barbatus and elucidates its effect on the soil seed bank in a semi-arid environment.     

Nest-plugging: interference competition in desert ants (Novomessor cockerelli and Pogonomyrmex barbatus)

Summary Pogonomyrmex barbatus and Novomessor cockerelli, sympatric species of harvester ants in the Lower Sonoran desert, compete for seed resources. This study reports on a method of interference competition. Early in the morning, before P. barbatus' activity period, N. cockerelli fills the nest entrances of P. barbatus with sand. This delays the beginning of the P. barbatus activity period for 1–3 h. P. barbatus colonies near N. cockerelli nests were more likely to be plugged. Nest-plugging shifts the typical daily sequence of P. barbatus activities, including the onset of foraging, forward towards midday, when high temperatures force the colony back inside the nest. P. barbatus colonies do not compensate for late emergence or events impeding foraging by increasing foraging rate. Thus nest-plugging by N. cockerelli decreases the foraging capacity of P. barbatus colonies.     

Harvester ant nests, soil biota and soil chemistry

Many ant species accumulate organic debris in the vicinity of their nests. These organic materials should provide a rich resource base for the soil biota. We examined the effect of harvester ant nests (Pogonomyrmex barbatus) on the soil community and soil chemistry. Ant nest soils supported 30-fold higher densities of microarthropods and 5-fold higher densities of protozoa than surrounding, control soils. The relative abundances of the major groups of protozoa differed as well: amoebae and ciliates were relatively overrepresented, and flagellates underrepresented, in ant nest versus control soils. Densities of bacteria and fungi were similar in the two soil types. Concentrations of nitrate, ammonium, phosphorus, and potassium were significantly higher in ant nest soils, while concentrations of magnesium, calcium, and water were similar in nest and control soils. Ant nest soils were marginally more acidic than controls. The results demonstrate that P. barbatus nests constitute a significant source of spatial heterogeneity in soil biota and soil chemistry in arid grasslands. Received: 17 March 1997 / Accepted: 10 June 1997     

Respiratory physiology and water relations of three species of Pogonomyrmex harvester ants (Hymenoptera: Formicidae)

The respiratory physiology and water relations of three harvester ant species (Pogonomyrmex rugosus Emery, P. occidentalis[Cresson] and P. californicus[Buckley]) were examined at three temperatures (15, 25 and 35°C) using a flow-through respirometry system. As intact ants tended to be active during testing, we performed a parallel set of experiments on individuals rendered motionless by decapitation. Both intact and decapitated ants exhibited discontinuous ventilation. Decapitation caused metabolic rate (V˙CO₂) and burst frequency to decrease in all three species. Burst volume either remained constant or increased after removal of the head, though mass-specific V˙CO₂ was unaffected except in P. rugosus. Mass-specific V˙CO₂s of headless harvesters were comparable with published values derived from motionless specimens of other ant species. The mean Q₁₀ for intact ants of all three species was 2.37; for decapitated insects the mean was 2.32. Respiratory water constituted a small (< 5%) fraction of total loss, and we believe that discontinuous ventilation does not act to conserve water in these organisms, although it may serve other functions.     

Effects of salinity on the establishment and early growth of Prosopis argentina and Prosopis alpataco seedlings in two contrasting soils: Implications for their ecological success

Abstract In arid zones, water availability is the most important factor limiting seedling establishment and plant distribution. However, within a region with a defined water regime, the physical and chemical properties of the soil could be the cause of the spatial pattern of plant communities. Prosopis argentina Burk. and Prosopis alpataco Phil. are sympatric at a regional scale but at the local scale they occupy clearly differentiated edaphic niches. Prosopis argentina reaches its ecological optimum in the sandy soils of active dunes, whereas P. alpataco achieves it in heavy, clayish, saline and periodically flooded soils. We studied the effect of salinity, and its interaction with soil type, on the establishment and early growth of these species in order to evaluate their adaptive mechanisms, and to analyse how this was related to the ecological success of these species. Salinity affected emergence and early growth of P. argentina and P. alpataco seedlings differently. Higher salinity led to decreased height, total biomass and shoot and root biomass of plants in both species but the effect was stronger in P. argentina than in P. alpataco, and greater in clayish than in sandy soils. These results would indicate that exclusion of P. argentina from clayish and saline soils would occur during emergence and the first stages of seedling establishment as a consequence of salinity. In P. alpataco other edaphic limitations, like texture or fertility in sandy soils, appear to be always more important. The osmotic effect evidenced by decreasing water and osmotic potentials of plants under saline conditions may be associated with growth reduction in both P. argentina and P. alpataco. However, this factor cannot explain differences in growth between species under salinity. The higher contents of Ca⁺⁺ and K⁺, and the lower contents of Na⁺ in P. alpataco suggest that the greater tolerance of this species depends on its ability to regulate and control absorption and transport of ions.     

Dietary similarity and foraging range of two seed-harvester ants during resource fluctuations

Summary Diets of desert seed-harvester ants Veromessor pergandei and Pogonomyrmex rugosus were studied for 3 years at two habitats where they are common and sympatric. Diets of the two species were similar, consisting mainly (87% of 23,913 seeds) of three annual plant species (Schismus arabicus, Plantago insularis, and Pectocarya recurvata). Diets converged following a drought in Winter/Spring 1984 which reduced seed production during this time. Foraging range of P. rugosus almost doubled following the drought while foraging range of V. pergandei remained constant. Neither ant species move their nests once established rendering them effectively sessile granivores. This limits the dietary options of a given colony and may explain interspecific dietary convergence.     

Bird effects on organic processes in soils from five microhabitats on a nunatak with and without breeding snow petrels in Dronning Maud Land, Antarctica

Five microhabitat types with varying degrees of bird influence were examined. Soils were collected from open polygons, under mosses and bird nests on a nunatak with breeding snow petrels (Pagodroma nivea) and from open polygons and under mosses on a non-bird nunatak. Nutrient levels (total N and P, nitrate, nitrite and ammonia), moisture levels and δ ¹⁵N values were determined and the organic processes of nitrogen fixation (acetylene reduction) and soil respiration (CO₂ flux) were examined. Nests represented the most favourable microhabitat type for soil respiration having the highest nutrient levels and most favourable temperature and moisture regimes. The soils under mosses were also favourable and appear to act as a nutrient sink for nutrients originating from the nests. The open polygons were the least favourable for biological activity. There was little nitrogen fixation in any of the soils except for the soils under mosses from the non-bird nunatak. Fixation is possibly limited in favourable microhabitat types on the bird nunatak by high nitrogen levels. These results were confirmed by the δ ¹⁵N results, which had high values typical of a seabird signal in the soils from the bird nunatak and values near zero, typical of soils containing fixed nitrogen, on the non-bird nunatak. Received: 3 March 1997 / Accepted: 30 March 1998     

Neighborhood density and reproductive potential in harvester ants

When neighbors compete for resources, the characteristics of a neighborhood may affect fitness. We examined the relationship between reproductive success and the density and size/age characteristics of neighbors in a population of the seed-eating ant, Pogonomyrmex barbatus, in which the ages of all colonies were known. Reproductive success was estimated by trapping and counting the number of alate, reproductive ants emerging from the nest for the annual mating flight. Alate production was negatively related to neighborhood density. Decreased production of alates by more crowded colonies may be due to competition for food with surrounding colonies. Neighbor size/age was unrelated to alate production. If alate production is correlated with lifetime reproductive success, these results suggest that selection favors colonies that monopolize more space, whatever the size of neighboring colonies. Received: 12 February 1996 / Accepted: 6 September 1996     

Soil temperature, digging behaviour, and the adaptive value of nest depth in South American species of Acromyrmex leaf-cutting ants

In leaf-cutting ants, workers are expected to excavate the nest at a soil depth that provides suitable temperatures, since the symbiotic fungus cultivated inside nest chambers is highly dependent on temperature for proper growth. We hypothesize that the different nesting habits observed in Acromyrmex leaf-cutting ants in the South American continent, i.e. superficial and subterranean nests, depend on the occurrence, across the soil profile, of the temperature range preferred by workers for digging. To test this hypothesis, we first explored whether the nesting habits in the genus Acromyrmex are correlated with the prevailing soil temperature regimes at the reported nest locations. Second, we experimentally investigated whether Acromyrmex workers engaged in digging use soil temperature as a cue to decide where to excavate the nest. A bibliographic survey of nesting habits of 21 South American Acromyrmex species indicated that nesting habits are correlated with the soil temperature regimes: the warmer the soil at the nesting site, the higher the number of species inhabiting subterranean nests, as compared to superficial nests. For those species showing nesting plasticity, subterranean nests occurred in hot soils, and superficial nests in cold ones. Experimental results indicated that Acromyrmex lundi workers use soil temperature as an orientation cue to decide where to start digging, and respond to rising and falling soil temperatures by moving to alternative digging places, or by stopping digging, respectively. The soil temperature range preferred for digging, between 20°C and maximally 30.6°C, matched the range at which colony growth would be maximized. It is suggested that temperature-sensitive digging guides digging workers towards their preferred range of soil temperature. Workers’ thermopreferences lead to a concentration of digging activity at the soil layers where the preferred range occurs, and therefore, to the construction of superficial nests in cold soils, and subterranean ones in hot soils. The adaptive value of the temperature-related nesting habits, and the temperature-sensitive digging, is further discussed.     

Rediscovery of the workerless inquiline antPogonomyrmex colei and additional notes on natural history (Hymenoptera: Formicidae)

Summary Pogonomyrmex colei is a workerless inquiline ant known only from nests ofP. rugosus, its closest relative. Ten of 776 (1.3%) host nests were parasitized at a site in central Arizona, while none of 1499 potential host colonies were parasitized at two other locales. Colonies ofP. colei are perennial, and host alate females in 9 of 10 colonies demonstrates that host queens survive parasitism. Three of 10 colonies died over 19 colony years of observation, while only 1 of 601 colonies became newly parasitized. Mating occurs in morning for up to 2–3 days following summer and fall rains and in afternoon during cool fall days. Mating is intranidal just outside the nest entrance, with males returning to the natal nest. MaleP. colei may be flightless because their wing area is reduced compared to host males. Females fly from the nest and locate potential host colonies by following trunk trails. Workers are the largest barrier to nest establishment, as they removed over 90% ofP. colei females placed in trunk trails or that entered host nests. Males and females ofP. colei andP. anergismus, the only other congeneric inquiline species, are diminutive compared to their hosts, with females 30% lighter than host workers. Fat content is lower and water content is higher inP. colei andP. anergismus females than in their hosts.     

Ecology and genetics of hybrid zones in the southern African Pycnonotus bulbul species complex

Lloyd, P., Craig, A.J.F.K., Hulley, P.E., Essop, M.F., Bloomer, P. & Crowe, T.M. 1997. Ecology and genetics of hybrid zones in the southern African Pycnonotus bulbul species complex. Ostrich 68 (2–4): 90–96.

The closely related Blackeyed Bulbul Pycnonotus barbatus, Cape Bulbul P. capensis and Redeyed Bulbul P. nigricans have parapatric to locally sympatric distributions within southern Africa. Extensive hybridization along narrow transition zones between each of the three species pairs is described in a region of the Eastern Cape province, South Africa. The transition zones coincide with ecotones between different vegetation types, which in turn follow escarpments or mountain ranges. The lack of population density depressions within the hybrid zones, together with the variability of the hybrids, suggests the hybrids are viable. Sharp step clines in various phenotypic characters are described across the P. barbatus/P. nigricans hybrid zone. A mtDNA analysis found evidence of possible introgression between P. barbatus and P. capensis. All eight P. barbatus x P. nigricans hybrids analysed possessed P. barbatus mtDNA, suggesting the existence of either positive assortative mating or strong directional selection, but our data are unable to distinguish which. Our results do not support the dynamic-equilibrium model, but are compatible with the bounded-hybrid-superiority model. We conclude that the maintenance of the parapatric distributions of the different taxa is due mainly to differences in environmentally-associated fitness between parental phenotypes or among parental and hybrid phenotypes along an ecotone, with the narrowness of the hybrid zones maintained by the steepness of the environmental gradients crossing them.     

Temporal patterns of seed use and availability in a guild of desert ants

ABSTRACT.

• 1 Temporal patterns of seed use were studied from late winter to autumn in three species of seed-harvesting ants in the Sonoran Desert. Measures of effective foraging activity, dietary niche breadth and dietary niche overlaps were obtained each month and were tested for correlation with estimates of the available seed resource.
• 2 Seeds were the only numerically important type of food in the diets of all species.
• 3 The ants partitioned the resource according to both seed species and seed size, although there was considerable overlap.
• 4 Pheidole xerophila had the smallest forager body size and is a specialist on small seeds because it harvested them in greater proportion than their rank in the soils and expanded its diet to larger seeds only when the abundance of small seeds declined.
• 5 When the abundance of the small seeds of Bouteloua barbata decreased, the middle-sized ant, Veromessor pergandei, showed a decrease in foraging activity, increase in niche breadth, and a decrease in overlap with P.xerophila.
• 6 Seed size preferences of V.pergandei did not vary seasonally, except that during the month of highest seed abundance, V.pergandei showed no size preference.
• 7 Pogonomyrmex rugosus was the largest ant; it preferred larger seeds and was inactive when small seeds were most abundant. Seasonal foraging activity and niche parameters were random in relation to seed abundance.
• 8 We suggest that nocturnal foraging by P.rugosus during the summer months was a response to interference with diurnal foraging by either predation frorn horned lizards or competition from V.pergandei.
• 9 Seasonal abundance of small seeds explains most of the seasonal foraging patterns of P.xerophila and V.pergandei. The summertime abundance of larger seeds during years of adequate precipitation may account for the seasonal activity patterns of P.rugosus.

     

Short-Term Vegetation and Soil Responses to Mechanical Destruction of Rabbit (Oryctolagus cuniculus L.) Warrens in an Australian Box Woodland

We studied the impact of disturbance by rabbits on plants and soils along a gradient out from the center of ripped rabbit warrens in an Australian semiarid woodland. Five years after the warrens were ripped, the impact of rabbits was still apparent. The cover of bare soil declined, and the cryptogam cover increased with increasing distance from the warren mound. However, litter cover, plant cover, and plant diversity remained unchanged with increasing distance from the mounds. Differences in plant composition were apparent with increasing distance from the mounds, with three species, Schismus barbatus, Salsola kali var. kali, and Chenopodium melanocarpum dominating the mounds, whereas the perennial grass Austrostipa scabra dominated the nonwarren control surfaces. Two species, Crassula sieberana and S. barbatus, dominated the active soil seed bank on ripped warrens. The mounds had the lowest number of species in the soil seed bank, whereas the warren edge microsite had the greatest. Ripped and unripped warrens differed substantially in their complement of species, and ripped warrens contained an order of magnitude fewer active warren entrances compared with unripped warrens. Ripped warrens also had significantly more plant cover than unripped warrens. Taken together, our results reinforce the view that rabbits have a destructive effect on surface soils and vegetation in semiarid woodlands and suggest that restoration of the original woodland vegetation after warren ripping is likely to be a slow and ongoing process.     

The process of population increase and patterns of resource utilization of two spider mites,Oligonychus ununguis (Jacobi) andPanonychus citri (McGregor), under experimental conditions (Acari: Tetranychidae)

Summary Pattern of population growth and characteristics of habitat utilization and of migration by two species of spider mites were studied under experimental conditions. The population growth ofOligonychus ununguis (Jacobi) on a chestnut occurred only on a single mite-release leaf over a long period, and few individuals moved away. Most of the 2nd progeny generation females of this species emigrated from the mite-release leaf as well as the sapling by means of ballooning threads. During this growth period, population density on the mite-release leaf levelled off, whereas that on the sapling increased. In contrast, the foundress ofPanonychus citri (McGregor) on citrus actively moved over several neighbouring leaves, and until the 2nd progeny generation females emerged, individuals were distributed over all the sapling leaves by means of walking. Emigration from the sapling was not observed until the 2nd progeny females emerged, and after that the mites emigrated by means of ballooning threads. The population density ofP. citri on the sapling levelled off and was rather decreased on the mite-release leaf at the time of mite emigration. Comparing the changing pattern of the relative degree of aggregation (m ^*/m) measured in two different units between these two species, the pattern ofm ^*/m in 1 cm² on the mite release leaf inO. ununguis resembled that of the unit of leaves on the sapling inP. citri. This result as well as behavioural observations indicate that migration ofO. ununguis is the movement from leaf to leaf and that ofP. citri from sapling to sapling. It is, therefore, concluded that the boundary of the microhabitat is a single leaf forO. ununguis but sapling or foliage forP. citri. This work was presented in Annual Meeting of Jap. Soc. Appl. Ent. Zool., 1981 in Okayama.     

Food robbing in ants,a form of interference competition

Summary Food robbing is a special form of interference competition in ants. It has been frequently observed in Myrmecocystus mimicus, which waylay returning foragers of several Pogonomyrmex species at their nests and take insect prey, particularly termites, away from them. Myrmecocystus more successfully robs prey from P. desertorum and P. maricopa than from P. barbatus. Usually only those Pogonomyrmex nests are affected by Myrmecocystus prey robbing that are not farther away than 10 m from the Myrmecocystus nest. Some M. mimicus workers show a high specialization in prey robbing. Other cases of food robbing in ants are described.     

Indirect effects of granivory by harvester ants: plant species composition and reproductive increase near ant nests

Summary Of 36 plant species surveyed, 6 were significantly associated with nests of the desert seed-harvester ant Veromessor pergandei or Pogonomyrmex rugosus; two other plant species were significantly absent from ant nests. Seeds of two common desert annuals, Schismus arabicus and Plantago insularis, realize a 15.6 and 6.5 fold increase (respectively) in number of fruits or seeds produced per plant growing in ant nest refuse piles compared to nearby controls. Mass of individual S. arabicus seed produced by plants growing in refuse piles also increased significantly. Schismus arabicus, P. insularis and other plants associated with ant nests do not have seeds with obvious appendages attractive to ants. Dispersal and reproductive increase of such seeds may represent a relatively primitive form of ant-plant dispersal devoid of seed morphological specializations. Alternatively, evolution of specialized seed structures for dispersal may be precluded by the assemblage of North American seed-harvester ants whose workers are significantly larger than those ants normally associated with elaiosome-attached seed dispersal. Large worker size may permit consumption of elaiosome and seed.     

Myrmica rubra microhabitat selection and putative ecological impact

1. Myrmica rubra (European fire ant) has invaded northern latitude coastal areas in North America. This macroscale distribution suggests that M. rubra is moisture‐ and temperature‐limited, but the distribution of the invaded range may reflect the legacy of original introduction locations preserved by limited dispersal. 2. This study examined a two‐decade population change in M. rubra (1994–2015) and the microscale abiotic (moisture and temperature), biotic (plants), anthropogenic (pesticide) and physiological (thermal tolerance) limits on the invasion at the Tifft Nature Preserve in Buffalo, NY (U.S.A.). Changes in the abundance of native ants and other invertebrates were also examined. 3. Despite localised declines with pesticide treatments, overall M. rubra forager abundance increased 27% between 1994 and 2015. Abundance increased the most in the warmest areas (native ants were unaffected by temperature), but M. rubra colony locations were strongly linked to higher soil moisture and lower soil temperature. Myrmica rubra ants also exhibited relatively low thermal tolerances consistent with high‐latitude and high‐elevation ants. 4. Where local M. rubra populations increased the most, native ant species decreased, and where local M. rubra populations declined, native ant species increased. Some arthropod species had lower abundance with M. rubra presence, but the impacts were less striking. 5. Myrmica rubra population growth was promoted at the microhabitat scale where relatively higher temperatures prompted foraging, and relatively lower temperatures and high moisture supported nesting. These results suggest that macroscale M. rubra invaded‐range distributions in northern climates near coastal areas are replicated at the microscale where the ant prefers cooler, moister microsites.