Wolf pack rendezvous site selection in Greece is mainly affected by anthropogenic landscape features

In wolves, most offspring mortality occurs within the first 6–8 months of their life. As wolf pups pass this entire period at either the den or rendezvous sites, their selection by wolf packs may affect pup survival and recruitment. Rendezvous sites are important for pup survival as they are used during summer and early autumn, when intense human activity may increase pup mortality. Adult wolves and pups can be killed by livestock guarding dogs during summer and intentionally or accidentally during large game hunting in autumn. This study describes factors related to rendezvous site selection in order to enhance their protection and management. We studied the rendezvous site selection of 30 wolf packs in central and northern Greece between 1998 and 2010, after locating 35 sites using the simulated howling survey method and telemetry. We considered a series of environmental and anthropogenic predictors of wolf rendezvous site selection at two spatial scales. At the landscape-population scale, wolves selected rendezvous sites below 1,200 m asl, with large inter-site distance (mean, 12.9 km), and avoided partially forested or open habitats, indicating preference for covered, spaced areas with seasonally stable resources. At the home range scale, wolves selected rendezvous sites away from forest roads and villages, close to water sources, and in areas with low forest fragmentation, indicating avoidance of human presence and disturbance. In the summer of 2011, we used an ensuing resource selection model (RSF, AUC?=?0.818) to successfully locate seven new rendezvous sites outside our previous survey area, verifying the utility of prediction maps (all new sites were at areas with 0.8–1 model probability). Rendezvous prediction maps can be used to reduce field effort when monitoring wolf populations, assess livestock predation risk, design protected areas, and reduce human disturbance on reproductive wolf packs.     

Wolf denning behaviour in response to external disturbances and implications for pup survival

We conducted a 15-year study (1999–2014) of wolf denning behavior in Naliboki Forest, Belarus, to determine the extent to which some external factors (rain, mosquitoes, human disturbances) and litter size are negatively correlated with pup survival until autumn.For this purpose we entirely documented denning areas of 30 breeding wolf pairs. Initial denning area includes the natal den site and (ii) secondary den sites between which the parent wolves transported sucessively the litter during the rearing season.The number of den sites per denning area ranged from 1 to 4. The distances in kilometres differed significantly between den sites. The total number of dens (surface dens and burrows) was higher in the natal den site compared with the other den sites. The number of surface dens per den site was higher compared with burrows in den sites 1, 2 and 3 but not den site 4. A total of 168 pups was found over the 15 years. Litter sizes ranged from 1 to 11 (average 5.6 ± 1.9). The rate of pup survival ranged from 0 to 100% and averaged 54.5 ± 29%.We examined the bivariate association between environmental conditions (mosquitoes, rainfall and human presence) and number of surface dens, total number of burrows and number of den sites. Environmental conditions were assessed from spring to mid-summer. Results show that there was a positive relationship between environmental conditions and number of den sites, and also that more difficult environmental conditions (rainy weather, many mosquitoes) decreased pup survival both in absolute and in relative terms. A cluster analysis showed that pup survival was also influenced by human presence. Wolves probably have a more complicated denning behaviour following stressful event which may lead to less effectiveness for feeding or protecting pups. The number of pups surviving was also found to be lower when there were many (8–10) pups because den shifting is more risky in this case. Burrows improve pup survival. Our results tend to suggest that, in our study area, a high mortality rate of wolf pups could be related to external and internal factors of stress.     

The Effects of Breeder Loss on Wolves

Abstract Managers of recovering wolf (Canis lupus) populations require knowledge regarding the potential impacts caused by the loss of territorial, breeding wolves when devising plans that aim to balance population goals with human concerns. Although ecologists have studied wolves extensively, we lack an understanding of this phenomenon as published records are sparse. Therefore, we pooled data (n = 134 cases) on 148 territorial breeding wolves (75 M and 73 F) from our research and published accounts to assess the impacts of breeder loss on wolf pup survival, reproduction, and territorial social groups. In 58 of 71 cases (84%), ≥1 pup survived, and the number or sex of remaining breeders (including multiple breeders) did not influence pup survival. Pups survived more frequently in groups of ≥6 wolves (90%) compared with smaller groups (68%). Auxiliary nonbreeders benefited pup survival, with pups surviving in 92% of cases where auxiliaries were present and 64% where they were absent. Logistic regression analysis indicated that the number of adult-sized wolves remaining after breeder loss, along with pup age, had the greatest influence on pup survival. Territorial wolves reproduced the following season in 47% of cases, and a greater proportion reproduced where one breeder had to be replaced (56%) versus cases where both breeders had to be replaced (9%). Group size was greater for wolves that reproduced the following season compared with those that did not reproduce. Large recolonizing (>75 wolves) and saturated wolf populations had similar times to breeder replacement and next reproduction, which was about half that for small recolonizing (≤75 wolves) populations. We found inverse relationships between recolonizing population size and time to breeder replacement (r= —0.37) and time to next reproduction (r= —0.36). Time to breeder replacement correlated strongly with time to next reproduction (r=0.97). Wolf social groups dissolved and abandoned their territories subsequent to breeder loss in 38% of cases. Where groups dissolved, wolves reestablished territories in 53% of cases, and neighboring wolves usurped territories in an additional 21% of cases. Fewer groups dissolved where breeders remained (26%) versus cases where breeders were absent (85%). Group size after breeder loss was smaller where groups dissolved versus cases where groups did not dissolve. To minimize negative impacts, we recommend that managers of recolonizing wolf populations limit lethal control to solitary individuals or territorial pairs where possible, because selective removal of pack members can be difficult. When reproductive packs are to be managed, we recommend that managers only remove wolves from reproductive packs when pups are ≥6 months old and packs contain ≥6 members (including ≥3 ad-sized wolves). Ideally, such packs should be close to neighboring packs and occur within larger (≥75 wolves) recolonizing populations.     

Den shifting by wolves in semi-wild landscapes in the Deccan Plateau, Maharashtra, India

Reproductive success is crucial for the survival and persistence of any species. The Deccan Biogeographic Zone of India is the stronghold of a population of the Indian wolf Canis lupus pallipes . Gaining a better understanding of the den-use pattern of wolves in different areas in this zone is thus vital for their conservation and management. The wolves excavated multiple dens in our study sites and kept shifting their litters among them. Major disturbance factors around denning sites were active stone quarries, traffic, crop harvesting and livestock movement. One wolf pack used 14 dens in four breeding seasons. Discriminant function analysis indicated that den shifting by wolves was not entirely governed by disturbance levels at den sites. Increasing age of pups was one of the main factors associated with den shifting rather than the magnitude of disturbance ( χ ²=34.26, d.f.=12, P <0.001). Tolerance to disturbance around dens during the early stages of pup development was negatively correlated ( r =−0.519, P <0.05) with availability of water and age of pups ( r =−0.613, P <0.01). Excavation of multiple dens by wolves was apparently related to den shifting, which seemingly is a survival strategy of wolves in these semi-wild human-dominated landscapes in the Deccan Biogeographic Zone. Principal components analysis indicated that during the initial stages of pup development, nature of the land, den orientation and distance of the den from roads were important cues in addition to age category of pups for den shifting. The analysis also suggested that factors such as distance of the new den where pups are to be transferred, distance from water source and availability of fox Vulpes bengalensis holes for den use were also important.     

Survival of captive and free-ranging Harlequin Ducks (Histrionicus histrionicus) following surgical liver biopsy

We measured intra- and postoperative mortality rates of captive and free-ranging Harlequin Ducks (Histrionicus histrionicus) undergoing surgical liver biopsy sampling for determination of the induction of cytochrome P4501A, a biomarker of oil exposure. Liver biopsies were taken from and radio transmitters were implanted into 157 free-ranging Harlequin Ducks over three winters (55 in 2000, 55 in 2001, and 47 in 2002). No birds died during surgery, but seven (4.5%) died during recovery from anesthesia (three in 2001 and four in 2002). None of the deaths could be attributed directly to the liver biopsy. Four of the 150 (2.7%) birds that were released died in the 2 wk period after surgery. All post-release deaths occurred in 2001; no birds died after release in 2000 or 2002. No mortalities of 36 captive birds occurred during surgery or recovery or in the 2 wk period following surgery. Hemorrhage was a minor problem with one captive bird. Surgical liver biopsies appear to be a safe procedure, but anesthetic complications may occur with overwintering ducks.     

Estimation of Successful Breeding Pairs for Wolves in the Northern Rocky Mountains,USA

Abstract: Under the Endangered Species Act, documenting recovery and federally mandated population levels of wolves (Canis lupus) in the Northern Rocky Mountains (NRM) requires monitoring wolf packs that successfully recruit young. United States Fish and Wildlife Service regulations define successful breeding pairs as packs estimated to contain an adult male and female, accompanied by ≥2 pups on 31 December of a given year. Monitoring successful breeding pairs will become more difficult following proposed delisting of NRM wolves; alternatives to historically intensive methods, appropriate to the different ecological and regulatory context following delisting, are required. Because pack size is easier to monitor than pack composition, we estimated probability a pack would contain a successful breeding pair based on its size for wolf populations inhabiting 6 areas in the NRM. We also evaluated the extent to which differences in demography of wolves and levels of human-caused mortality among the areas influenced the probability of packs of different sizes would contain successful breeding pairs. Probability curves differed among analysis areas, depending primarily on levels of human-caused mortality, secondarily on annual population growth rate, and little on annual population density. Probabilities that packs contained successful breeding pairs were more uniformly distributed across pack sizes in areas with low levels of human mortality and stable populations. Large packs in areas with high levels of human-caused mortality and high annual growth rates had relatively high probabilities of containing breeding pairs whereas those for small packs were relatively low. Our approach can be used by managers to estimate number of successful breeding pairs in a population where number of packs and their sizes are known. Following delisting of NRM wolves, human-caused mortality is likely to increase, resulting in more small packs with low probabilities of containing breeding pairs. Differing contributions of packs to wolf population growth based on their size suggests monitoring successful breeding pairs will provide more accurate insights into population dynamics of wolves than will monitoring number of packs or individuals only.     

Direct Estimation of Early Survival and Movements in Eastern Wolf Pups

Abstract: Determining juvenile survival and recruitment rates is essential to assess status and viability of animal populations. Currently, the demographic attributes of juvenile carnivores, specifically wolves (Canis lycaon), are poorly known but of considerable conservation interest. We measured survival and dispersal rates for 51 juvenile (age 3.5–31 weeks) wolves in Algonquin Provincial Park, Canada, from 2004 to 2005, using implantable very high frequency transmitters. Monthly pup survival was high (0.970, 95% CI = 0.951–0.990) and constant from June to November, and most pup mortality was from natural causes. Pups dispersed as early as age 15 weeks, and monthly dispersal rates were high for young pups (min. = 0.008, 95% CI = 0.000–0.019; max. = 0.030, 95% CI = 0.010–0.050). We failed to detect any influence of pack or litter size on pup survival or probability of dispersal. Radiotelemetry offers an individual-based monitoring technique capable of providing direct assessment of wolf pup survival and movements, with rigorous estimation of survival and dispersal rates and quantification of cause-specific mortality.     

Surveying Predicted Rendezvous Sites to Monitor Gray Wolf Populations

ABSTRACT We used rendezvous site locations of wolf (Canis lupus) packs recorded during 1996–2006 to build a predictive model of gray wolf rendezvous site habitat in Idaho, USA. Variables in our best model included green leaf biomass (Normalized Difference Vegetation Index), surface roughness, and profile curvature, indicating that wolves consistently used wet meadow complexes for rendezvous sites. We then used this predictive model to stratify habitat and guide survey efforts designed to document wolf pack distribution and fecundity in 4 study areas in Idaho. We detected all 15 wolf packs (32 wolf pack-yr) and 20 out of 27 (74%) litters of pups by surveying <11% of the total study area. In addition, we were able to obtain detailed observations on wolf packs (e.g., hair and scat samples) once we located their rendezvous sites. Given an expected decrease in the ability of managers to maintain radiocollar contact with all of the wolf packs in the northern Rocky Mountains, rendezvous sites predicted by our model can be the starting point and foundation for targeted sampling and future wolf population monitoring surveys.     

ATYPICAL PUP REARING STRATEGIES BY SEA OTTERS

Eight tagged sea otter (Enhydra lutris) pups in central Prince William Sound, Alaska, weighed 6–15 kg at the time of separation from their mother. Four pups weighing 15 kg were able to forage successfully on their own. Three pups weighing ≤9 kg had negligible chances of survival and apparently were abandoned by sick females. Abandonment of a pup may reduce the burden on a sick female, enabling recovery and subsequent reproduction. One of the three sick females that abandoned a pup in this study recovered and pupped again. Abandonment of pups should occur most often in populations where females are stressed by poor food resources. Reassociation with a previous offspring, as observed once in this study, also may occur most frequently in food-limited populations where reproductive failures are most common and pup survivorship is significantly increased by additional maternal assistance.     

Gray wolf density and its association with weights and hematology of pups from 1970 to 1988.

We examined weights and hematologic profiles of gray wolf (Canis lupus) pups and the associated wolf density in the east-central Superior National Forest of northeastern Minnesota (USA) during 1970 to 1988. We collected weight and hematologic data from 117 pups (57 females, 60 males) during 1 September to 22 November each year. The wolf density (wolves/800 km2) trend was divided into three phases: high (72 +/- 7), 1970 to 1975; medium (44 +/- 2), 1976 to 1983; and low (27 +/- 2), 1984 to 1988. Wolf numbers declined (P = 0.0001) 39 and 63% from 1970 to 1975 to 1976 to 1983 and from 1970 to 1975 to 1984 to 1988, respectively. Weight was similar between male and female pups and did not vary as wolf density changed. Mean hemoglobin (P = 0.04), red (P = 0.0001) and white blood cells (P = 0.002), mean corpuscular volume, mean corpuscular hemoglobin concentration and mean corpuscular hemoglobin (P = 0.0001) did differ among the multi-annual phases of changing wolf density. Weight and hematologic data also were compared to values from captive wolf pups. The high, but declining wolf density was associated with macrocytic, normochromic anemia in wolf pups, whereas the lowest density coincided with a hypochromic anemia. Although hematologic values show promise for assessing wolf pup condition and wolf population status, they must be used cautiously until data are available from other populations.     

Cloning endangered gray wolves (Canis lupus) from somatic cells collected postmortem

The objective of the present study was to investigate whether nuclear transfer of postmortem wolf somatic cells into enucleated dog oocytes, is a feasible method to produce a cloned wolf. In vivo-matured oocytes (from domestic dogs) were enucleated and fused with somatic cells derived from culture of tissue obtained from a male gray wolf 6h after death. The reconstructed embryos were activated and transferred into the oviducts of naturally synchronous domestic bitches. Overall, 372 reconstructed embryos were transferred to 17 recipient dogs; four recipients (23.5%) were confirmed pregnant (ultrasonographically) 23-25 d after embryo transfer. One recipient spontaneously delivered two dead pups and three recipients delivered, by cesarean section, four cloned wolf pups, weighing 450, 190, 300, and 490g, respectively. The pup that weighed 190g died within 12h after birth. The six cloned wolf pups were genetically identical to the donor wolf, and their mitochondrial DNA originated from the oocyte donors. The three live wolf pups had a normal wolf karyotype (78, XY), and the amount of telomeric DNA, assessed by quantitative fluorescence in situ hybridization, was similar to, or lower than, that of the nuclear donor. In conclusion, the present study demonstrated the successful cloning of an endangered male gray wolf via interspecies transfer of somatic cells, isolated postmortem from a wolf, and transferred into enucleated dog oocytes. Therefore, somatic cell nuclear transfer has potential for preservation of canine species in extreme situations, including sudden death.     

How does parental role influence the activity and movements of breeding wolves?

We studied the activity and movements of a male and a female wolf during breeding in the Bieszczady Mountains, Poland. The female was less active and mobile and on average closer to the den than the male. The male was less active and mobile when the pack size was seven compared to a year when the pack consisted only of the breeding pair. We conclude that the roles played by breeding males and females rearing pups influence their activity, and that breeding males in larger packs move less during the nursing period because of help by other pack members.     

Dog Pups’ Attractiveness to Humans Peaks at Weaning Age

The domestic dog (Canis lupus familiaris) is characterized by greatly reduced parenting investment compared with the wild type wolf (C. l. lupus) from which it is descended. Unlike wolf pups, which are reared by both parents into their second year of life, dog pups are abandoned by their mother at weaning around eight weeks of age. This relatively small parental involvement may contribute to the high pup mortality observed in dogs not living as pets. We hypothesized that people would find dog pups most attractive around weaning age when conspecific parental care is significantly reduced and pup mortality rate is high. Younger and older pups would benefit less from human intervention because in the former case the mother is providing care, and in the latter their survival is already compromised. To test this hypothesis, 51 participants rated the attractiveness of 39 black and white headshot photographs presented on a computer screen of dog pups from three breeds (Jack Russell Terrier, Cane Corso, and White Shepherd), from birth to 7 months old. In line with our hypothesis, attractiveness of Cane Corsos peaked at 6.3 weeks of age; Jack Russell Terriers’ attractiveness peaked at 7.7 weeks; and White Shepherds were most attractive at 8.3 weeks. There were also differences in attractiveness between the breeds, with Cane Corsos rated less attractive than the other two breeds. If this attractiveness motivates humans to care for the dog pups and thereby improves pup survival, this could confer significant advantages to dogs, and may contribute to our understanding of the process of domestication.     

Some Aspects of the Population Ecology of Wolves, Alaska

Information on Alaskan wolf populations was obtained from examinationof bounty records, 4,150 wolf radii and ulnae, 1,262 wolf carcasses,and from observations of wolves inhabiting an area of 20,000square miles where wolves were protected. Pregnant adult female wolves averaged 6.5 fetuses; two-year-oldfemales averaged 5.3 fetuses;female pups were not sexually mature. In Alaska, wolves conceive from late February through earlyApril but most females breed in March. Multiparous females breedearlier than first breeders. Multiparous females produce anaverage of 7.3 ova and 6.5 implanted fetuses. The loss of ovafrom ovulation to implantation is significant. Multiparous femalesproduce more ova than first breeders; the difference is highlysignificant. Mortality of pups rather than the lack of initial productionof pups is believed to be the reason for the observed variationsin the proportion of pups in wolf populations. Wolf packs includemembers in all categories of sex and age during the breedingseason. Size of the pack is an indicator of abundance. Wolves in an area where they were protected increased at anaverage rate of 20–30% per year during an 11-year period.     

Response of Wolves to Experimental Disturbance at Homesites

ABSTRACT Events during the denning period (parturition to first autumn) often determine the reproductive success of wolves (Canis lupus). Consequently, there is concern about the potential adverse effects of human-caused disturbance at wolf den and rendezvous sites (homesites), but relatively little information on this subject is available. We conducted standardized experimental disturbance treatments at 12 unique wolf homesites in the Northwest Territories, Canada, during summers 2002 and 2003. The treatment consisted of an intruder approaching a homesite once per day for 3 consecutive days and recording behavioral responses, response distance, and response intensity of wolves. We counted pups and estimated their ages prior to the initial treatment at each site. Adult wolves moved pups at 3 of 6 treated homesites in each year. The amount and type of known human activity within a pack's home range did not influence whether adults moved pups in response to the treatment. The response intensity of wolves to the treatment was inversely related to the amount of human activity near a homesite. There was no relationship between the distance at which wolves responded to the intruder and the amount or type of human activity. There was a positive relationship between increasing age of pups and their relocation in response to the treatment. Reproductive success was not influenced by the treatment or by the amount and type of human activity. Treated sites were used by wolves the following year in the same proportion as untreated sites. It appears that pups are most vulnerable early in the year when less mobile; therefore, managers should consider age of pups before human activity at or near wolf homesites occurs.     

First report of captive New Guinea dingo (Canis dingo hallstromi) den-digging and parental behavior

New Guinea dingoes (NGDs) (Canis dingo hallstromi; Troughton [1957] Proc Roy Soc new South Wells 1955–1956:93–94) have been kept in zoos since 1956. Almost nothing is known of their wild behavior. These observations of a captive pair are the first documentation of natal den‐digging and parental behavior for this taxon. The main den, excavated near the top of a 1.5 m hill, consisted of a rounded chamber about 50.8 cm deep, with an entrance about 30.5 cm high and 40.6 cm wide. The dam frequently moved the pups from the natal den to secondary locations for short periods during the day and then back to the den, starting when the pups were 2 weeks old. When the pups were between 5 and 12 weeks of age, both parents regularly regurgitated for them. The sire expressed escalating threat behavior toward the male pup starting when the pup was 5 months old, and the female began threatening the female pups at about 6 months of age. Rejection of same‐sex offspring is usual for captive NGDs as the next breeding season approaches. Zoo Biol 30:445–450, 2011. © 2010 Wiley‐Liss, Inc.     

Neonatal anesthesia for studies of hamster parental behavior when infanticidal aggression is a possibility

Experiments involving investigation of the neuroendocrine basis for paternal care in rodents risk activation of aggressive behavior toward pups. To minimize pain and suffering during tests of parental responsiveness requiring retrieval of a displaced pup to its nest, a method of anesthetizing the pup was developed in Djungarian hamsters, Phodopus campbelli. A surgical plane of anesthesia, as measured by criteria, such as respiratory depression, loss of the pedal reflex, and failure to increase respiratory rate or to vocalize in response to handling, was achieved by use of intraperitoneal administration of a combination of ketamine and xylazine. Both parents (tested separately) expressed normal behavior toward anesthetized pups. In random order, a saline-injected or anesthetized pup was displaced from its nest in the home cage. There were no differences in pick-up or retrieval rates between saline and anesthetized pups for either parent. A third test using an unmanipulated pup confirmed that parental behavior was not reduced toward an anesthetized pup. However, if anesthetized pups were tested first among littermates, retrieval by males was less likely. This method will, therefore, underestimate retrieval behavior in males, but not females. Adult male hamsters that had never been parents also expressed expected behavior by attacking the pup in 45% of cases. This method provides an efficient and effective means of protecting pups while allowing adults to express a wide range of parental and infanticidal behaviors. It also has application in behavioral screening of transgenic strains toward unrelated young.     

Inbreeding is reduced by female-biased dispersal and mating behavior in Ethiopian wolves

Molecular tools have enabled wildlife researchers to obtainaccurate information on the kinship, mating behavior, and dispersalof individuals. We genotyped 192 Ethiopian wolves (n = 29 packs)in the Bale Mountains for 17 microsatellite loci to 1) elucidatekinship within and between packs, 2) assess parentage of pups,and 3) evaluate whether inbreeding is avoided by dispersal and/ormating behavior. Mean pairwise relatedness within packs (R =0.39) was significantly greater than that estimated from randomassignment of individuals to packs. However, breeding pairswere most often unrelated, suggesting that female-biased dispersalreduces inbreeding. We assigned maternity to 49 pups and paternityto 47 pups (n = 12 litters) using a combination of exclusion,likelihood analyses (using CERVUS software), and sibship reconstruction.Multiple paternity occurred in 33% of litters; extrapack paternityaccounted for 28% of all resolved paternities, occurring in50% of litters. We found no evidence that extrapack copulationsreduce inbreeding; however, more detailed analyses may elucidatethe effect of recent population declines and demographic disturbancesdue to recurring disease outbreaks. The adaptive advantagesof female-biased dispersal and the observed mating system arediscussed in relation to Ethiopian wolf sociobiology and ecology.     

Analgesic effects of tramadol, carprofen or multimodal analgesia in rats undergoing ventral laparotomy

In this study, the authors evaluated the analgesic efficacy of tramadol (an opioid-like analgesic), carprofen (a nonsteroidal anti-inflammatory drug) and a combination of both drugs (multimodal therapy) in a rat laparotomy model. The authors randomly assigned rats to undergo either surgery (abdominal laparotomy with visceral manipulation and anesthesia) or anesthesia only. Rats in each group were treated with tramadol (12.5 mg per kg body weight), carprofen (5 mg per kg body weight), a combination of tramadol and carprofen (12.5 mg per kg body weight and 5 mg per kg body weight, respectively) or saline (anesthesia control group only; 5 mg per kg body weight). The authors administered analgesia 10 min before anesthesia, 4 h after surgery or (for the rats that received anesthesia only) anesthesia and 24 h after surgery or anesthesia. They measured locomotor activity, running wheel activity, feed and water consumption, body weight and fecal corticosterone concentration of each animal before and after surgery. Clinical observations were made after surgery or anesthesia to evaluate signs of pain and distress. The authors found that carprofen, tramadol and a combination of carprofen and tramadol were all acceptable analgesia regimens for a rat laparotomy model.     

Maternal responsiveness following differential pup treatment and mother-pup interactions

Pituitary-adrenal and behavioral responsiveness to rat pups which had been handled or shocked was assessed in mothers at Days 2, 9, and 16 postpartum. Mothers were exposed to these pups either directly where mother-infant contact was permitted or indirectly (pups placed in a wire mesh basket) where mother-infant contact was prevented. Handled and shocked pups elicited differential pituitary-adrenal responsiveness when mother-infant contact was prevented following pup treatment. Mothers responded more to shocked than to handled pups. While pup treatments elicited different patterns of maternal behavior, direct mother-infant contact subsequent to pup treatment eliminated the differential pattern of pituitary-adrenal responsiveness. These findings were discussed in terms of a parallel between behavioral and pituitary-adrenal indices of maternal responsiveness and the proposed role of maternal mediation in early experience effects.