Kris J. Hundertmark

Reproductive Characteristics of Alaskan Moose

Many aspects of moose (Alces alces) reproduction are poorly documented. Therefore, we quantified the estrous cycle, estrous length, gestation period, fetal development, and birth mass of calves. We evaluated empirical relationships among maternal age, mass, and previous breeding parity with litter size and neonatal sex ratio. Increased attentiveness by the bull signaled estrous behavior in cows. Estrous females did not increase activity. The estrous cycle varied from 22 to 28 days (x = 24.4 days) and did not lengthen with each successive cycle. The cycle of primiparous females was shorter (P = 0.05) than pluriparous females. Gestation averaged 231 days (SD = 5.4 days) and did not differ (P > 0.05) between primiparous and pluriparous females, litters of 1 or 2 calves, among 5 years of study, or between cows bred their first or second overt estrus

MITOCHONDRIAL PHYLOGEOGRAPHY OF MOOSE (ALCES ALCES) IN NORTH AMERICA

Abstract Nucleotide variation was assessed from the mitochondrial control region of North American moose (Alces alces) to test predictions of a model of range expansion by stepping-stone dispersal and to determine whether patterns of genetic variation support the current recognition of 4 subspecies. Haplotypes formed a star phylogeny indicative of a recent expansion of populations. Values of nucleotide and haplotype diversity were low continent-wide but were greatest in the central part of the continent and lowest in peripheral populations. Despite low mitochondrial diversity, moose exhibited a high degree of differentiation regionally, which was not explained by isolation by distance. Our data indicate a pattern of colonization consistent with a large central population that supplied founders to peripheral populations (other than Alaska), perhaps through rare, long-distance dispersal events (leptokurtic dispersal) rather than mass dispersal by a stepping-stone model. The colonization scenario does not account for the low haplotype diversity observed in Alaska, which may be derived from a postcolonization bottleneck. Establishment of peripheral populations by leptokurtic dispersal and subsequent local adaptation may have been sufficient for development of morphological differentiation among extant subspecies.

Fecal progestagen concentration as an indicator of the estrous cycle and pregnancy in moose

We established a noninvasive technique for monitoring the estrous cycle and pregnancy in moose (Alces alces). We collected fecal samples daily from nonpregnant yearling females (n = 3) for the duration of 3-4 estrous cycles, and weekly throughout gestation from 3 pregnant adult cows. We analyzed samples by radioimmunoassay (RIA) to quantify the concentration of progestagen metabolites. Peak luteal phase concentrations of progestagens were 4.5 ± 0.9 (SD) μg/g compared with 0.5 ± 0.1 μg/g during the follicular phase. Observed (n = 7) and suspected (n = 2) matings all occurred within ±2 days of the nadir in progestagen excretion. Fecal progestagens were above peak luteal phase concentrations by the eighth week of gestation (>7 μg/g of feces, P < 0.001), permitting accurate pregnancy detection by this time. Relative concentrations of fecal progestagens during the follicular phase, luteal phase, and pregnancy were 1:10:35, respectively. Monitoring fecal progestagen is a useful noninvasive technique for tracking ovarian activity and pregnancy in moose.

Estimating Abundance of Sitka Black-Tailed Deer Using DNA From Fecal Pellets

ABSTRACT Densely vegetated environments have hindered collection of basic population parameters on forest-dwelling ungulates. Our objective was to develop a mark—recapture technique that used DNA from fecal pellets to overcome constraints associated with estimating abundance of ungulates in landscapes where direct observation is difficult. We tested our technique on Sitka black-tailed deer (Odocoileus hemionus sitkensis) in the temperate coastal rainforest of Southeast Alaska. During 2006–2008, we sampled fecal pellets of deer along trail transects in 3 intensively logged watersheds on Prince of Wales Island, Alaska. We extracted DNA from the surface of fecal pellets and used microsatellite markers to identify individual deer. With genotypes of individual deer, we estimated abundance of deer with moderate precision (±20%) using mark—recapture models. Combining all study sites, we identified a 30% (SE = 5.1%) decline in abundance during our 3-year study, which we attributed to 3 consecutive severe winters. We determined that deer densities in managed land logged >30 years ago (7 deer/km2, SE = 1.3) supported fewer deer compared to both managed land logged <30 years ago (10 deer/km2, SE = 1.5) and unmanaged land (12 deer/km2, SE = 1.4). Our study provides the first estimates of abundance (based on individually identified deer) for Sitka black-tailed deer and the first estimates of abundance of an unenclosed ungulate population using DNA from fecal pellets. Our tool enables managers to accurately and precisely estimate the abundance of deer in densely vegetated habitats using a non-invasive approach.

Population Structure and Genetic Diversity of Moose in Alaska

Moose (Alces alces) are highly mobile mammals that occur across arboreal regions of North America, Europe, and Asia. Alaskan moose (Alces alces gigas) range across much of Alaska and are primary herbivore consumers, exerting a prominent influence on ecosystem structure and functioning. Increased knowledge gained from population genetics provides insights into their population dynamics, history, and dispersal of these unique large herbivores and can aid in conservation efforts. We examined the genetic diversity and population structure of moose (n = 141) with 8 polymorphic microsatellites from 6 regions spanning much of Alaska. Expected heterozygosity was moderate (H(E) = 0.483-0.612), and private alleles ranged from 0 to 6. Both F(ST) and R(ST) indicated significant population structure (P < 0.001) with F(ST) < 0.109 and R(ST) < 0.125. Results of analyses from STRUCTURE indicated 2 prominent population groups, a mix of moose from the Yakutat and Tetlin regions versus all other moose, with slight substructure observed among the second population. Estimates of dispersal differed between analytical approaches, indicating a high level of historical or current gene flow. Mantel tests indicated that isolation-by-distance partially explained observed structure among moose populations (R(2) = 0.45, P < 0.01). Finally, there was no evidence of bottlenecks either at the population level or overall. We conclude that weak population structure occurs among moose in Alaska with population expansion from interior Alaska westward toward the coast.

Two reciprocally monophyletic mtDNA lineages elucidate the taxonomic status of Mountain gazelles (Gazella gazella)

Mountain gazelles (Gazella gazella) rank among the most critically endangered mammals on the Arabian Peninsula. Past conservation efforts have been plagued by confusion about the phylogenetic relationship among various ‘phenotypically discernable’ populations, and even the question of species boundaries was far from being certain. This lack of knowledge has had a direct impact on conservation measures, especially ex situ breeding programmes, hampering the assignment of captive stocks to potential conservation units. Here, we provide a phylogenetic framework, based on the analysis of mtDNA sequences (360 bp cytochrome b and 213 bp Control Region) of 126 individuals collected from the wild throughout the Arabian Peninsula and from captive stocks. Our analyses revealed two reciprocally monophyletic genetic lineages within the presumed species Gazella gazella: one ‘northern clade’ on the Golan Heights (Israel/Syrian border) and one genetically diverse larger clade from the rest of the Arabian Peninsula including the Arava Valley (Negev, Israel). Applying the Strict Phylogenetic Species Concept (sensu Mishler & Theriot, 2000) allows assigning species status to these two major clades.

Dead before detection: addressing the effects of left truncation on survival estimation and ecological inference for neonates

Summary 1. Neonate survival is a key life history trait, yet remains challenging to measure in wild populations because neonates can be difficult to capture at birth. Estimates of survival from neonates that are opportunistically captured might be inaccurate because some individuals die before sampling, resulting in data that are left truncated. The resulting overestimation of survival rates can further affect ecological inference through biased estimates of covariate effects in survival models, yet is not addressed in most studies of animal survival. Here, we quantify the effects of left truncation on survival estimates and subsequent ecological inference. 2. Vaginal implant transmitters (VITs) enable capture of ungulates at birth, yielding data without left truncation. The effects of left truncation on survival estimation were quantified using age-dependent survival models for VIT and opportunistically captured neonatal deer. Differences in daily survival rates (DSRs) and cumulative survival probability were calculated for the first 70 days of life. In addition, left truncation was simulated by removing fawns that died during the first 1 or 2 days of life from the VIT-caught sample, isolating the effect of left truncation. 3. Cumulative probability of survival during the first 70 days of life was overestimated by 7–23% for fawns caught opportunistically compared with those caught by VIT, depending on model design. Differences in DSRs were large at age 1 day, but had converged by age 30 days. Simulated left truncation resulted in overestimates of survival of up to 31%. Model selection and covariate coefficients were strongly affected by left truncation, producing spurious ecological inference, including changes to sign and/or magnitude of inferred effects of all covariates. 4. We recommend (i) every effort be made to capture neonates; (ii) consistent capture methods, using at least in part non-truncating techniques, be implemented across years and study areas; and (iii) exclusion of left-truncated data from survival estimates until DSRs converge with those calculated from non-truncated data. This work emphasizes the importance of accounting for left truncation in survival estimation for any species with strong age-dependent survival in order to prevent biased conclusions produced by sampling method rather than true ecological effects.

Phylogenetic analysis of mitochondrial DNA sequences reveals polyphyly in the goitred gazelle (Gazella subgutturosa)

Goitred gazelle (Gazella subgutturosa) rank among the most endangered mammals on the Arabian Peninsula and the Asian steppes. Past conservation efforts have been plagued by confusion about the phylogenetic relationship among various—phenotypically discernable—populations, and even the question of species boundaries was far from being certain. This lack of knowledge had a direct impact on conservation measures, especially ex situ breeding programmes, hampering the assignment of captive stocks to potential conservation units. Here, we provide a phylogenetic framework, based on the analysis of mtDNA sequences of a number of individuals collected from the wild and captivity throughout the species’ natural range. Our analyses revealed a polyphyly within the presumed species of G. subgutturosa resulting in two distinct clades: one on the Arabian Peninsula, Iraq, Jordan, Syria and Turkey (Gazella marica; sand gazelle) and one genetically diverse larger clade from the rest of its Asian range (G. subgutturosa; goitred gazelle). Additionally, we provide a quick method (PCR-RFLP) to analyse the taxonomic affiliation of captive gazelles that will be used for re-introductions into the wild.

Mountain Goat Survival in Coastal Alaska: Effects of Age, Sex, and Climate

ABSTRACT Ecological theory predicts that individual survival should vary between sex and age categories due to differences in allocation of nutritional resources for growth and reproductive activities. During periods of environmental stress, such relationships may be exacerbated, and affect sex and age classes differently. We evaluated support for hypotheses about the relative roles of sex, age, and winter and summer climate on the probability of mountain goat (Oreamnos americanus) survival in coastal Alaska. Specifically, we used known-fates analyses (Program MARK) to model the effects of age, sex, and climatic variation on survival using data collected from 279 radio-marked mountain goats (118 M, 161 F) in 9 separate study areas during 1977–2008. Models including age, sex, winter snowfall, and average daily summer temperature (during Jul–Aug) best explained variation in survival probability of mountain goats. Specifically, our findings revealed that old animals (9+ yr) have lower survival than younger animals. In addition, males tended to have lower survival than females, though differences only existed among prime-aged adult (5–8 yr) and old (9+ yr) age classes. Winter climate exerted the strongest effects on mountain goat survival; summer climate, however, was significant and principally influenced survival during the following winter via indirect effects. Furthermore, old animals were more sensitive to the effects of winter conditions than young or prime-aged animals. These findings detail how climate interacts with sex and age characteristics to affect mountain goat survival. Critically, we provide baseline survival rate statistics across various age, sex, and climate scenarios. These data will assist conservation and management of mountain goats by enabling detailed, model-based demographic forecasting of human and/or climate-based population impacts.

Individual identification of Sitka black-tailed deer (Odocoileus hemionus sitkensis) using DNA from fecal pellets

We tested a protocol for extracting DNA from fecal pellets from Sitka black-tailed deer (Odocoileus hemionus sitkensis) and evaluated genotyping performance of previously developed microsatellite markers as well as a suite of new markers designed specifically for this study. We screened 30 microsatellites, and identified 7 (23%) loci including 4 new markers, that fit well into a single multiplex and consistently genotyped deer with low error rates. DNA was extracted from 2,408 fecal-pellet samples. Of those, 1,240 (52%) were genotyped successfully at all 7 markers allowing identification of 634 genetically unique deer. Using DNA from fecal pellets collected in the field was an effective technique for identifying and distinguishing among deer.