Was Aldo Leopold Right about the Kaibab Deer Herd?

In ecology textbooks prior to the 1970s, Aldo Leopold’s classic story of predator control, overpopulation of deer, and habitat degradation on the Kaibab Plateau during the 1920s epitomized predator regulation of herbivore populations. However, the story disappeared from texts in the late 20th century after several papers noted uncertainties in estimations of the deer population and provided alternative explanations. We re-examined the case study by determining the age structure of aspen (Populus tremuloides Michx.) on the plateau. Aspen comprises the majority of deer browse in the summer, and the absence of a normal cohort of aspen from the 1920s would indicate deer overpopulation. The number of aspen (at 1.4 m) dating to the 1920s was an order of magnitude lower than the null expectation. Other periods of unusual numbers of aspen included high numbers of aspen dating to the 1880s and 1890s (when regular surface fires ceased), few aspen dating from 1953 to 1962 (after a second irruption of the deer population), and very high numbers from 1968 to 1992 (coincident with widespread logging). These convergent lines of evidence support the idea of extreme deer herbivory in the 1920s, consistent with food limitation of deer at high populations (bottom–up control) and predation limitation at low deer populations (top–down control). Some uncertainty remains within the overall story, and this level of ambiguity is common in case studies that involve population ecology, land management, and people at the scale of 1,000 km² and 100 years. A complete version of the Kaibab deer story and its history would be a valuable, realistic case study for ecology texts.     

Adaptive Lévy Walks in Foraging Fallow Deer

Background

Lévy flights are random walks, the step lengths of which come from probability distributions with heavy power-law tails, such that clusters of short steps are connected by rare long steps. Lévy walks maximise search efficiency of mobile foragers. Recently, several studies raised some concerns about the reliability of the statistical analysis used in previous analyses. Further, it is unclear whether Lévy walks represent adaptive strategies or emergent properties determined by the interaction between foragers and resource distribution. Thus two fundamental questions still need to be addressed: the presence of Lévy walks in the wild and whether or not they represent a form of adaptive behaviour.

Methodology/Principal Findings

We studied 235 paths of solitary and clustered (i.e. foraging in group) fallow deer (Dama dama), exploiting the same pasture. We used maximum likelihood estimation for discriminating between a power-tailed distribution and the exponential alternative and rank/frequency plots to discriminate between Lévy walks and composite Brownian walks. We showed that solitary deer perform Lévy searches, while clustered animals did not adopt that strategy.

Conclusion/Significance

Our demonstration of the presence of Lévy walks is, at our knowledge, the first available which adopts up-to-date statistical methodologies in a terrestrial mammal. Comparing solitary and clustered deer, we concluded that the Lévy walks of solitary deer represent an adaptation maximising encounter rates with forage resources and not an epiphenomenon induced by a peculiar food distribution.     

Deer antler – A novel model for studying organ regeneration in mammals

Deer antler is the only mammalian organ that can fully grow back once lost from its pedicle – the base from which it grows. Therefore, antlers probably offer the most pertinent model for studying organ regeneration in mammals. This paper reviews our current understanding of the mechanisms underlying regeneration of antlers, and provides insights into the possible use for human regenerative medicine. Based on the definition, antler renewal belongs to a special type of regeneration termed epimorphic. However, histological examination failed to detect dedifferentiation of any cell type on the pedicle stump and the formation of a blastema, which are hallmark features of classic epimorphic regeneration. Instead, antler regeneration is achieved through the recruitment, proliferation and differentiation of the single cell type in the pedicle periosteum (PP). The PP cells are the direct derivatives of cells resident in the antlerogenic periosteum (AP), a tissue that exists in prepubertal deer calves and can induce ectopic antler formation when transplanted elsewhere on the deer body. Both the AP and PP cells express key embryonic stem cell markers and can be induced to differentiate into multiple cell lineages in vitro and, therefore, they are termed antler stem cells, and antler regeneration is a stem cell-based epimorphic regeneration. Comparisons between the healing process on the stumps from an amputated mouse limb and early regeneration of antlers suggest that the stump of a mouse limb cannot regenerate because of the limited potential of periosteal cells in long bones to proliferate. If we can impart a greater potential of these periosteal cells to proliferate, we might at least be able to partially regenerate limbs lost from humans. Taken together, a greater understanding of the mechanisms that regulate the regeneration of antlers may provide a valuable insight to aid the field of regenerative medicine.This article is part of a Directed Issue entitled: Regenerative Medicine: the challenge of translation.     

Lack of Mother—Offspring Relationships in White-Tailed Deer Capture Groups

ABSTRACT Behavioral studies of white-tailed deer (Odocoileus virginianus) often assign mother-offspring relationships based on common capture of juveniles with adult deer, assuming that fawns associate closely with mothers. We tested this assumption using genetic parentage to assess mother-offspring relationships within capture groups based on data from 10 polymorphic microsatellite loci. At the 80% confidence level, we assigned maternity to 43% and 51% of juveniles captured with an adult female in 2 respective study areas. Capture with their mother did not differ by sex of juveniles in either study area, and limiting our analysis to capture groups that most represent family groups (i.e., one ad F with 1–3 juv) did not increase maternity assignment (35%). Our results indicate that common capture may be a poor indicator of mother-offspring relationships in many field settings. We recommend genetic verification of family relationships.     

Mother to Offspring Transmission of Chronic Wasting Disease in Reeves’ Muntjac Deer

The horizontal transmission of prion diseases has been well characterized in bovine spongiform encephalopathy (BSE), chronic wasting disease (CWD) of deer and elk and scrapie of sheep, and has been regarded as the primary mode of transmission. Few studies have monitored the possibility of vertical transmission occurring within an infected mother during pregnancy. To study the potential for and pathway of vertical transmission of CWD in the native cervid species, we used a small cervid model–the polyestrous breeding, indoor maintainable, Reeves’ muntjac deer–and determined that the susceptibility and pathogenesis of CWD in these deer reproduce that in native mule and white-tailed deer. Moreover, we demonstrate here that CWD prions are transmitted from doe to fawn. Maternal CWD infection also appears to result in lower percentage of live birth offspring. In addition, evolving evidence from protein misfolding cyclic amplification (PMCA) assays on fetal tissues suggest that covert prion infection occurs in utero. Overall, our findings demonstrate that transmission of prions from mother to offspring can occur, and may be underestimated for all prion diseases.     

Genetics of Hemoglobin in the Deer Mouse, PEROMYSCUS MANICULATUS . I. Multiple α- and β-Globin Structural Loci

Genetic data, together with molecular structure studies, have demonstrated that the complex hemoglobin phenotypes in four subspecies of P. maniculatus are generated by at least four, and probably five, globin structural loci. The Hba and Hbc loci apparently arose by duplication of an ancestral α-type locus, while Hbb, Hbd, and Hbe apparently were derived from a β-type locus. The Hba and Hbb structural loci are electrophoretically monomorphic, while Hbc, Hbd, and Hbe are each polymorphic for at least two electrophoretic alleles. Alleles at Hbc and Hbd segregate independently. The globin products of Hbc and Hbd are electrophoretically indistinguishable; therefore, it is impossible to enumerate true gene frequencies in population surveys. Combined evidence from Peromyscus, Mus and Rattus indicates a remarkable similarity in the numbers of duplicated globin structural loci and in their linkage relationships to coat color loci.     

Control of Ixodes scapularis and Amblyomma americanum Through use of the ‘4-poster’ Treatment Device on Deer in Maryland

Deer self-treatment devices (‘4-posters’) were evaluated for their efficacy in reducing populations of blacklegged ticks, Ixodes scapularis, and lone star ticks, Amblyomma americanum. At each of three locations in Maryland, 25 ‘4-posters’ were operated in study areas of approximately 5.18 km². Populations of host-seeking ticks were monitored by flagging of treated areas and similar untreated control areas without ‘4-posters.’ From 1998 to 2002 the percent mortalities achieved were 69, 75.8 and 80 at the three study sites infested with I. scapularis nymphs, and 99.5 and 95.3 for A. americanum nymphs at the two sites where this species occurred. This revised version was published online in July 2006 with corrections to the Cover Date.     

Ranging Movements of Female Roe Deer: Do Home-loving Does Roam to Mate?

Bimonthly changes in home range size of seven mature roe deer Capreolus capreolus L. females were analysed over one year. Great stability of home range size and high site fidelity was common to all females, which remained bimonthly within areas of 6.1–20.8 ha. Two groups of females could be identified on the basis of bimonthly variations in their home range sizes: females with larger areas (LHR females), who decreased their ranges in the birth season only, and females with smaller home ranges (SHR females), who increased significantly their home ranges during the rut. The most likely explanation is that the areas inhabited by SHR females included high-quality food resources allowing survival even in small home ranges. In the rut, SHR females may have expanded their ranges to increase mating opportunities.     

Do Père David's Deer Lose Memories of Their Ancestral Predators?

Whether prey retains antipredator behavior after a long period of predator relaxation is an important question in predator-prey evolution. Père David''s deer have been raised in enclosures for more than 1200 years and this isolation provides an opportunity to study whether Père David''s deer still respond to the cues of their ancestral predators or to novel predators. We played back the sounds of crows (familiar sound) and domestic dogs (familiar non-predators), of tigers and wolves (ancestral predators), and of lions (potential naïve predator) to Père David''s deer in paddocks, and blank sounds to the control group, and videoed the behavior of the deer during the experiment. We also showed life-size photo models of dog, leopard, bear, tiger, wolf, and lion to the deer and video taped their responses after seeing these models. Père David''s deer stared at and approached the hidden loudspeaker when they heard the roars of tiger or lion. The deer listened to tiger roars longer, approached to tiger roars more and spent more time staring at the tiger model. The stags were also found to forage less in the trials of tiger roars than that of other sound playbacks. Additionally, it took longer for the deer to restore their normal behavior after they heard tiger roars, which was longer than that after the trial of other sound playbacks. Moreover, the deer were only found to walk away after hearing the sounds of tiger and wolf. Therefore, the tiger was probably the main predator for Père David''s deer in ancient time. Our study implies that Père David''s deer still retain the memories of the acoustic and visual cues of their ancestral predators in spite of the long term isolation from natural habitat.     

Modern Dynamics of the Red Deer (<Emphasis Type="Italic">Сervus elaphus maral</Emphasis>) Population in the Caucasian State Nature Reserve

The dynamics of 19 local red deer (Сervus elaphus maral Ogilbi 1840) groupings in the Caucasian State Nature Biosphere Reserve in the period of 1996?2015 has been researched; the groupings were divided into three groups by their size: large, average, and small ones. It was found that the population decline at the end of 20th century has resulted in levelling of local deer groupings: all of them became small but none disappeared completely. Further population growth has resulted in the formation of large and average local groupings in central parts of the nature reserve; a gradual restoration of optimal ratios between those can be observed. In some peripheral areas of the biosphere reserve, the red deer population is not recovering, while the average size of the small groupings is decreasing. At least four local deer groupings have lost their independent status in the metapopulation structure of the species due to anthropogenic factors and continue their existence owing to neighboring local groupings on the basis of the “source?sink” system operation principle.     

Is Difference in Body Weight,Antler Length,Age or Dominance Rank related to the Number of Fights between Fallow Deer (Dama dama)?

Competitive interactions between mature male fallow deer were investigated to determine whether antler length, body weight, age or dominance rank were related to the number of fights between individuals. Four different hypotheses were tested; the first predicted that body weight and antler length indicate individual quality and, therefore, as the difference between competitors in body weight and antler length increases, there should be a corresponding decrease in fight rate. The second and third hypotheses predicted that, as difference in dominance rank increased there would be a decrease in fight rate between males either, as a risk reduction measure or because of inhibitory control by dominant males. A fourth hypothesis predicted that, where dominance rank does not mediate fight rate, that similarity between contestants based on age might be important. If this is the case, then as the difference in age between competitors increased there should be a decrease in fight rate. Our results show that when dominance rank is controlled for, there was no relation between body weight, antler length or age with fight rate. There was a negative relation between dominance rank difference and fight rate, a result that supports both the risk reduction and inhibitory hypotheses. There was an increasing tendency to fight with closely ranked males as cohorts reached peak reproductive age; males aged 5 yr fought with other 5‐yr‐old males based on rank difference and males aged 6 yr fought with other males aged six and across all age groups based on rank difference. This trend was not observed in 4 or 7‐yr‐old males. Our results suggest that males in prime breeding condition limit potential costs of fighting (such as time and energy) by only interacting with other males of similar rank.     

Technological Analysis of the World’s Earliest Shamanic Costume: A Multi-Scalar,Experimental Study of a Red Deer Headdress from the Early Holocene Site of Star Carr,North Yorkshire,UK

Shamanic belief systems represent the first form of religious practice visible within the global archaeological record. Here we report on the earliest known evidence of shamanic costume: modified red deer crania headdresses from the Early Holocene site of Star Carr (c. 11 kya). More than 90% of the examples from prehistoric Europe come from this one site, establishing it as a place of outstanding shamanistic/cosmological significance. Our work, involving a programme of experimental replication, analysis of macroscopic traces, organic residue analysis and 3D image acquisition, metrology and visualisation, represents the first attempt to understand the manufacturing processes used to create these artefacts. The results produced were unexpected—rather than being carefully crafted objects, elements of their production can only be described as expedient.     

Enemy release or invasional meltdown? Deer preference for exotic and native trees on Isla Victoria,Argentina

Abstract How interactions between exotic species affect invasion impact is a fundamental issue on both theoretical and applied grounds. Exotics can facilitate establishment and invasion of other exotics (invasional meltdown) or they can restrict them by re‐establishing natural population control (as predicted by the enemy‐release hypothesis). We studied forest invasion on an Argentinean island where 43 species of Pinaceae, including 60% of the world's recorded invasive Pinaceae, were introduced c. 1920 but where few species are colonizing pristine areas. In this area two species of Palearctic deer, natural enemies of most Pinaceae, were introduced 80 years ago. Expecting deer to help to control the exotics, we conducted a cafeteria experiment to assess deer preferences among the two dominant native species (a conifer, Austrocedrus chilensis, and a broadleaf, Nothofagus dombeyi) and two widely introduced exotic tree species (Pseudotsuga menziesii and Pinus ponderosa). Deer browsed much more intensively on native species than on exotic conifers, in terms of number of individuals attacked and degree of browsing. Deer preference for natives could potentially facilitate invasion by exotic pines. However, we hypothesize that the low rates of invasion currently observed can result at least partly from high densities of exotic deer, which, despite their preference for natives, can prevent establishment of both native and exotic trees. Other factors, not mutually exclusive, could produce the observed pattern. Our results underscore the difficulty of predicting how one introduced species will effect impact of another one.     

Do Red Deer Stags (Cervus elaphus) Use Roar Fundamental Frequency (F0) to Assess Rivals?

It is well established that in humans, male voices are disproportionately lower pitched than female voices, and recent studies suggest that this dimorphism in fundamental frequency (F0) results from both intrasexual (male competition) and intersexual (female mate choice) selection for lower pitched voices in men. However, comparative investigations indicate that sexual dimorphism in F0 is not universal in terrestrial mammals. In the highly polygynous and sexually dimorphic Scottish red deer Cervus elaphus scoticus, more successful males give sexually-selected calls (roars) with higher minimum F0s, suggesting that high, rather than low F0s advertise quality in this subspecies. While playback experiments demonstrated that oestrous females prefer higher pitched roars, the potential role of roar F0 in male competition remains untested. Here we examined the response of rutting red deer stags to playbacks of re-synthesized male roars with different median F0s. Our results show that stags’ responses (latencies and durations of attention, vocal and approach responses) were not affected by the F0 of the roar. This suggests that intrasexual selection is unlikely to strongly influence the evolution of roar F0 in Scottish red deer stags, and illustrates how the F0 of terrestrial mammal vocal sexual signals may be subject to different selection pressures across species. Further investigations on species characterized by different F0 profiles are needed to provide a comparative background for evolutionary interpretations of sex differences in mammalian vocalizations.     

The annual habitat selection of released Père David’s Deer in Dafeng Milu National Nature Reserve

The number of Père David’s (Elaphurus davidinus) deer which have been released into the wild is over one hundred in the third core area of Dafeng Milu National Nature Reserve. Annual habitat selection of released Père David’s deer was studied from November 2008 to November 2009. The results of our research showed that: in autumn and winter released Père David’s deer preferred to choose the locations at Spartina alterniflora community with higher plant density, higher plant height and larger aboveground biomass; in spring and summer released Père David’s deer preferred to choose the locations at S. alterniflora community with lower plant height, smaller aboveground biomass. Released Père David’s deer preferred to choose the locations at S. alterniflora community which are close to water sources and shallow bogs but have a moderate distance from the sea. Principal components analysis was performed on 8 ecological factors. The cumulative contribution of the first 3 principal components reached to 75% and these factors could represent the annual habitat features of released Père David’s deer. According to the results of principal component analysis, the influential factors for annual habitat selection of released Père David’s deer were classified separately as food factor (aboveground biomass, plant height), water factor (distance to water, distance from the shallow bogs), disturbance factor (distance from human disturbance).     

The annual habitat selection of released Père David’s Deer in Dafeng Milu National Nature Reserve

The number of Père David’s (Elaphurus davidinus) deer which have been released into the wild is over one hundred in the third core area of Dafeng Milu National Nature Reserve. Annual habitat selection of released Père David’s deer was studied from November 2008 to November 2009. The results of our research showed that: in autumn and winter released Père David’s deer preferred to choose the locations at Spartina alterniflora community with higher plant density, higher plant height and larger aboveground biomass; in spring and summer released Père David’s deer preferred to choose the locations at S. alterniflora community with lower plant height, smaller aboveground biomass. Released Père David’s deer preferred to choose the locations at S. alterniflora community which are close to water sources and shallow bogs but have a moderate distance from the sea. Principal components analysis was performed on 8 ecological factors. The cumulative contribution of the first 3 principal components reached to 75% and these factors could represent the annual habitat features of released Père David’s deer. According to the results of principal component analysis, the influential factors for annual habitat selection of released Père David’s deer were classified separately as food factor (aboveground biomass, plant height), water factor (distance to water, distance from the shallow bogs), disturbance factor (distance from human disturbance).     

Variability of Female Responses to Conspecific vs. Heterospecific Male Mating Calls in Polygynous Deer: An Open Door to Hybridization?

Males of all polygynous deer species (Cervinae) give conspicuous calls during the reproductive season. The extreme interspecific diversity that characterizes these vocalizations suggests that they play a strong role in species discrimination. However, interbreeding between several species of Cervinae indicates permeable interspecific reproductive barriers. This study examines the contribution of vocal behavior to female species discrimination and mating preferences in two closely related polygynous deer species known to hybridize in the wild after introductions. Specifically, we investigate the reaction of estrous female red deer (Cervus elaphus) to playbacks of red deer vs. sika deer (Cervus nippon) male mating calls, with the prediction that females will prefer conspecific calls. While on average female red deer preferred male red deer roars, two out of twenty females spent more time in close proximity to the speaker broadcasting male sika deer moans. We suggest that this absence of strict vocal preference for species-specific mating calls may contribute to the permeability of pre-zygotic reproductive barriers observed between these species. Our results also highlight the importance of examining inter-individual variation when studying the role of female preferences in species discrimination and intraspecific mate selection.     

Did the evolution of the phytoplankton fuel the diversification of the marine biosphere?

We discuss the possible links between the fossil record of marine biodiversity, nutrient availability and primary productivity. The parallelism of the fossil records of marine phytoplankton and faunal biodiversity implicates the quantity (primary productivity) and quality (stoichiometry) of phytoplankton as being critical to the diversification of the marine biosphere through the Phanerozoic. The relatively subdued marine biodiversity of the Palaeozoic corresponds to a time of relatively low macronutrient availability and poor food quality of the phytoplankton as opposed to the diversification of the Modern Fauna through the Mesozoic–Cenozoic. Increasing nutrient runoff to the oceans through the Phanerozoic resulted from orogeny, the emplacement of Large Igneous Provinces (LIPs), the evolution of deep-rooting forests and the appearance of more easily decomposable terrestrial organic matter that enhanced weathering. Positive feedback by bioturbation of an expanding benthos played a critical role in evolving biogeochemical cycles by linking the oxidation of dead organic matter and the recycling of nutrients back to the water column where they could be re-utilized. We assess our conclusions against a recently published biogeochemical model for geological time-scales. Major peaks of marine diversity often occur near rising or peak fluxes of silica, phosphorus and dissolved reactive oceanic phosphorus; either major or minor ⁸⁷Sr/⁸⁶Sr peaks; and frequently in the vicinity of major (Circum-Atlantic Magmatic Province) and minor volcanic events, some of which are associated with Oceanic Anoxic Events. These processes appear to be scale-dependent in that they lie on a continuum between biodiversification on macroevolutionary scales of geological time and mass extinction.     

Imperfect Batesian mimicry—the effects of the frequency and the distastefulness of the model

Batesian mimicry is the resemblance between unpalatable models and palatable mimics. The widely accepted idea is that the frequency and the unprofitability of the model are crucial for the introduction of a Batesian mimic into the prey population. However, experimental evidence is limited and furthermore, previous studies have considered mainly perfect mimicry (automimicry). We investigated imperfect Batesian mimicry by varying the frequency of an aposematic model at two levels of distastefulness. The predator encountered prey in a random order, one prey item at a time. The prey were thus presented realistically in a sequential way. Great tits (Parus major) were used as predators. This experiment, with a novel signal, supports the idea that Batesian mimics gain most when the models outnumber them. The mortalities of the mimics as well as the models were significantly dependent on the frequency of the model. Both prey types survived better the fewer mimics there were confusing the predator. There were also indications that the degree of distastefulness of the model had an effect on the survival of the Batesian mimic: the models survived significantly better the more distasteful they were. The experiment supports the most classical predictions in the theories of the origin and maintenance of Batesian mimicry.