Address correspondence to H. C. Rosenbaum, Wildlife Conservation Society, Science Resource Center, 2300 Southern Blvd., Bronx, NY 10460, or e-mail: hrosenbaum
Journal of Heredity, Volume 93, Issue 6, November 2002, Pperiods 389–399, https://doi.org/10.1093/jhered/93.6.389
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Abstract

To research whether demographic and also life-background traits are associated with genetic structure, we contrasted mtDNA lineperiods of individual humpearlier whales (Megaptera novaeangliae) via sighting and reproductive backgrounds of female humpago whales in between 1979 and 1995. Maternal lineage haploforms were acquired for 323 whales, either from direct sequencing of the mtDNA manage area (n = 159) or inferred from known relationships alengthy matrilines from the sequenced sample of individuals (n = 164). Sequence variation in the 550 bp of the control area characterized a total of 19 maternal family tree haploforms that formed 2 primary clades. Fecundity boosted significantly over the study period among females of several lineeras among the two clades. Individual maternal lineages and also various other clades were identified by considerable variation in fecundity. The detected heterogeneity of refertile success has actually the potential to substantially influence the frequency and also circulation of maternal lineages found in this population over time. Tright here were considerable yat an early stage results on adult resighting rates and calf survivorship based on examicountry of sighting backgrounds via differing capture-recapture probability models. These outcomes show that populace structure can be affected by interactions or associations in between reabundant success, genetic framework, and eco-friendly components in a organic population of long-lived mammals.

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Variation in reproductive success among people of either sex, and among grslrfc.orgs or cohorts, is famed in some wild populations of animals (Clutton-Brock 1988, 1989). It has generally been welcomed that heritcapability of life-history attributes regarded fitness, such as female fecundity, is low (Charlesworth 1994; Nunney 1995; Roff and Mousseau 1987). Differential mating success and also fecundity has actually been more freshly linked with individual variation in phenokind, age-structured effects, and also short-lived environmental readjust pertained to climate, population density, or habitat top quality (Clutton-Brock 1988; Hewichild 1997; Magrath 1989; Price and also Grant 1984; Sinervo and Zamudio 2001; Stoleboy and Beissinger 1997). While yearly variation in reproductive success has been argued to account for differences in oboffered heterozygosity and also better inreproduction coefficients (Scribner et al. 1993), there are few examples where heritability (genetic or cultural) is correlated through variable refertile success (Austerlitz and also Heyer 1998). Recent modeling argues that variation in reabundant success may impact the level of genetic diversity (Tiedemann and Milinkovitch 1999). More often, such corconnections through life-history traits are attributed to environmental, or even more extensively, stochastic distinctions.

It has lengthy been an objective of population genes to combine life-background parameters right into knowledge transforms in allele frequency (Wright 1951). However before, the lack of resolution of molecular markers and a paucity of appropriately detailed demographic framework and also life-history indevelopment have made this evaluation tough in nonhuman populaces (Clutton-Brock 1988, 1989). Only freshly have molecular markers permitted the straight research of the interactivity between populace genetic structure and mating units in a wider evolutionary context beyond the individual level. Correlation of trends in fecundity, survivorship, and also resighting for both ancestry and unique genetic lineeras within a natural population provides a new approach to a fundamental question of standard population genetics. Here we research the interactivity in between genetic framework and also fecundity in a long-lived nonhumale mammal and also the results any type of association can have on maternal lineage distribution and also vicapability in the study population.

The North Atlantic Ocean populace of humpback whales is composed of a number of fairly discrete feeding subpopulaces (Katona and Beard 1990) via matrilineal fidelity (Clapham and also Mayo 1987, 1990). One of these subpopulations is discovered in the Gulf of Maine throughout the months of April via December in the waters about Stellwagen Bank and also Jeffrey"s Ledge (Clapham et al. 1993; Weinaffluent et al. 1997) (Figure 1). This seasonally residential subpopulace of humpback whales has been stupassed away broadly given that the 1970s. Since people are identifiable by organic marmajesties on the ventral surchallenge of their tail flukes and dorsal fins (Katona and Whitehead 1981), fads of social company and also seasonal presence for Gulf of Maine humpago whales have been tracked from 1979 until the existing (e.g., Clapham 1992; Clapham and Mayo 1987; Weinrich 1991, 1998; Weinrich and also Kuhlberg 1991; Weinwealthy et al. 1997).

Mitochondrial DNA (mtDNA) as a maternal molecular marker has prstove to be beneficial in identifying populace structure and also confirming regular matrilineal recruitment of calves right into the Gulf of Maine from a widespread reproduction location (Baker et al. 1990; Clapham et al. 1993; Palsbøll et al. 1995). While populace structure shows up to be governed by maternal fidelity to a feeding area (Baker et al. 1990, 1994a; Palsbøll et al. 1995), the local circulation of whales throughout the Gulf of Maine appears mostly dependent on prey circulation and abundance (Payne et al. 1990; Weinaffluent et al. 1997) via minimal effects because of predators (Clapham 1996). It has actually been presumed that prey availcapability results in occasional disturbances in tempdental and also spatial circulation rather than considerable short-lived changes in genetic variation or survivorship of animals within the Gulf of Maine.

Dramatic changes in frequency and age structure of individual maternal lineperiods across time have actually likewise been oboffered wbelow considerable development, recirculation, and regional mass mortality have developed over the last three years (Baker et al. 1994b). The potential for combining molecular and also eco-friendly information to evaluate genetic results of natural occasions was demonstrated by Baker et al. (1994b). In that research, 8 of 10 victims of a organic mortality shared 2 of the much less common maternal family tree haplokinds in this area, offering the initially proof for association in between maternal family tree complace and demography.

In order to examine the potential interenergetic impacts of population genetic framework via survival and refertile success, we surveyed variations and also corconnections of life-history and also demographic parameters with genetic structure utilizing genealogical rebuilding of maternal lineperiods of humpback whales in the Gulf of Maine over a 16-year period (1979–1995). This approach was undertaken to determine if differential fecundity and refertile success are correlated with the evolution of maternal family tree or distinctive matrilines within this population. Pronounced variation in the circulation and frequency of maternal haplokinds within this population can aclimb from a connection in between maternal family tree varicapability and also fecundity, elevating questions about the procedure which perhaps web links the 2 and just how hereditary variation is construed for conservation and monitoring of an endangered species.

Materials and Methods

Sighting File and Capture-Recapture Histories

Extensive life-background information (spatial, tempdental, social, refertile, and also behavioral) have actually been accumulated for the majority of the sampled individuals and also their well-known relatives. Daily surveys were conducted (weather permitting) aboard commercial whale-watching watercrafts and dedicated vessels mainly operating from Gloucester, Massachusetts, from April 1 to November 1 in each year from 1979 to 1995. The examine location consisted of Stellwagen Bank and also Jeffrey"s Ledge, 2 underwater glacial deposits off the New England also coastline (Figure 1). Humpago whales were photographically figured out by pigment variation on the ventral surface of the tail flukes and also scarring and shape of the dorsal fin (Katona and also Whitehead 1981). Mother/offspring relationships were figured out by welcomed field techniques (Clapham and Mayo 1987, 1990; Weinrich et al. 1992). More details on field techniques have the right to be uncovered in Weinaffluent and Kuhlberg (1991) and also Weinwell-off et al. (1991).

Skin samples for genetic evaluation were collected utilizing a biopsy darting procedure (Lambertsen 1987; Weinrich et al. 1991) from 159 individually established free-ranging humpago whales in the southerly Gulf of Maine between 1988 and also 1993. Any other whales known to be part of the exact same matriline of a sampled whale via photographic identification research studies (e.g., calves of a sampled female, a mommy of a sampled calf, and the calf"s half-siblings) were assigned the very same mtDNA haploform. As our photographic identification work includes a maximum of three generations within a matriline, we feel the assumption that tbelow were no mutations in the mtDNA in between direct descendants is justified for the study.

Molecular Studies

Total genomic DNA was isolated from the epidermal layer of tworry biopsies utilizing standard proteinase K/phenol/chloroform measures (Sambrook et al. 1989). A percentage of the mtDNA regulate region, a noncoding area that is extremely variable in many type of vertebrates (Vigilant et al. 1991), was amplified making use of polymerase chain reaction (PCR). Two oligonucleotide primers (Dlp 1.5 and Dlp 5) have actually been emerged to amplify a 550 bp fragment within the mtDNA manage area of humpearlier whales (Baker et al. 1993) and from various other mammals (Kocher et al. 1989; Southern et al. 1988). This 550 bp area contains the majority of variable nucleotide positions in the mtDNA manage area of the humpback whale (Baker and also Medrano-Gonzalez, 2002; Baker et al. 1993). PCR commodities were purified utilizing Bio 101 Gene Clean II (hands-on sequencing) or Centrsymbol 100 columns (automated sequencing) according to the manufacturer"s protocol.

Automated DNA sequencing of PCR assets was done using cycle sequencing via fluorescently labeled dideoxy terminators and also an Applied Biosystems 373A DNA sequencer. Determination or confirmation of sex (from area monitorings of visibility or absence of a hemispherical lobe on the ventral surchallenge posterior to the genital slit or by association via a calf) was accomplished utilizing molecular methods. Gender was assigned by PCR amplification of a Y chromosome-particular sequence and also amplification and subsequent TaqI digestion of homologous regions on the X and also Y chromosomes (Palsbøll et al. 1992; Rictough et al. 1994).

Data Analysis

DNA sequence variation trends were defined into existing haplotype meanings for this species according to the nomenclature of Baker et al. (1993). Sequences for this percentage of the mtDNA regulate region were maintained for each individual in MacClade (version 3.01; Maddikid and Maddichild 1992). Alignment of DNA sequences from the mtDNA control area was perdeveloped utilizing MALIGN (Wheeler and also Gladstein 1992). The ancestry of Gulf of Maine mtDNA regulate area haplotypes was generated through parsimony utilizing both the branch and bound and also stepwise addition heuristic search choices in PAUP (version 4.0; Swofford 1998) and also rooted with homologous sequence from very closely connected intra- and interspecific outgrslrfc.orgs (Baker et al. 1993). Strict consensus trees were constructed from all trees saved in the branch and also bound search. Bootstrap values (Felsenstein 1985) were produced from 100 heuristic search replicates in PAUP (variation 4.0; Swofford 1998). Bremer supports (Bremer 1994) were produced through AUTODECAY (Erikschild 1998) utilizing 5 random-enhancement heuristic searcs for each node.

Fecundity Amongst Maternal Lineages

Correlations among maternal family tree haplokinds and also life history were tested using summary procedures such as fecundity, yearly survival, and recapture backgrounds. Survival data and reproductive propapers were sorted by maternal family tree haplotype for recognized females over the 16-year research duration. We calculated fecundity as the propercent of years in which a reproductively mature female was recorded via a calf in the time of the examine duration. This allowed us to assess the level to which individual maternal lineeras vary in reproductive success, while managing for sampling effort and also probcapacity of recapture. Heterogeneity in fecundity was examined based on resolution in the ancestry at various levels (I–V) consisting of 2 main lineperiods, individual lineperiods where sufficient data exist for statistical power, and also other clades listed below the 2 major clades where sufficient fecundity information and normally refixed grslrfc.orgings in the genealogy exist (view Figure 3 for a graphical depiction of these levels). These levels represent the miscellaneous ordered nodes in the ancestry. The genealogical analysis readily available independent justification to partition or pool rarer lineeras based upon levels of refixed nodes or branch assistance in order to test for corconnections with life background. Testing for associations of fecundity through maternal family tree at various genealogical levels enabled us to think about the persistence of any detected connection throughout the ancestry. Statistical meaning of distinctions between refertile success and family tree grslrfc.orgings (levels I–V) was figured out by utilizing a Kruskal–Wallis and also a Fisher"s precise test (two-tailed), owing to low (much less than five) counts in certain cells of a contingency table analysis. Statistical trial and error of all individual maternal lineeras (listed below level I) was not correct because of low numbers of females for particular lineperiods. To boost the power of the comparichild, fecundity was classified right into rates of much less than 0.25 (low), 0.25–0.49 (medium), or 0.5–1.0 (high), and also contrasted via the assorted levels throughout the genealogy with a Fisher"s specific test (two-tailed).

Temporal trends in fecundity were examined throughout the 16-year (1979–1995) research duration. Median fecundity was calculated among all females in aggregateways of well-known refertile age and also regressed, using least squares linear regression, against year of monitoring. Trends were examined among females of the 2 major clades (BCD versus IJK, level V). Documents were again also thin to contrast individual maternal lineage haploforms, yet fecundity was calculated for each female and contrasted among 4 added different maternal family tree combicountries based on resolution in the ancestry and where adequate data existed (level I: individual haplotypes C, D, I; level II: C, D1-D, I-I1-I2, K2-K3, and J-J1-J2-J3; level III: BCD, I-I1-I2, K1A-K2-K3-J-J1-J2-J3-J4; and also level IV: C, D-D1, and I-I1-I2, J-J1-J2-J3) for a finer resolution analysis (Figure 3). Multiple tests in this research were carried out in a nested fashion and also were not repeated tests of the same hypothesis on exactly the same datasets. The latter is the typical case where a correction for multiple comparisons is required (Perneger 1998).

Because individual females were observed for variable durations (1–16 years), approximates of calving prices were potentially vulnerable to sampling error based upon the variety of years a female was observed. For example, females via a higher number of resightings are likely to be oboffered with a calf more generally than females through fewer resightings. In order to eliminate this predisposition, the probability of nonfreedom in between recapture and refertile backgrounds was investigated. Price quotes of the coeffective of variation (CV) in refertile rates came to be a lot even more precise (CV 2; df = 36; χ2 = 40.64; P =.273). Tright here was likewise no substantial connection in between the variety of years a female was observed and also her reproductive rate (df = 86; r = 0.0355; P =.745).

Reabundant Success and Annual Survival

Survival rates were based upon resighting data of individually identified whales in the Gulf of Maine from 1979 with 1995. Due to the fact that some individuals were not seen in some years, approximates of survival based on monitorings might confound survival via the likelihood of resighting (Clobert et al. 1987; Cooch et al. 1996). We therefore employed maximum likelihood methods to obtain sepaprice approximates of survival (ϕ) and also resighting rates (p), making use of the software application SURGE 4.2 (Cooch et al. 1996).

Capture-recapture models incorporate three presumptions (1) ϕ and also p are homogeneous within any kind of subgrslrfc.org, (2) marking is precise and irreversible, and (3) the resighting or fatality of any kind of individual is independent of the fate of other people (Burnham et al. 1987). The recommfinished method for experimentation assumption 1 is to use the goodness-of-fit tests (tests 2 and also 3) of the regimen RELEASE (Burnham et al. 1987; Cooch et al. 1996). However, our data were as well sparse for this evaluation. Instead, we thought about calves and also adults individually, and also supplied an as a whole goodness-of-fit chi-square test to compare observed frequencies with version predictions. For the second assumption over, individuals were figured out by herbal and permanent variations in ventral fluke pigmentation (Katona and also Whitehead 1981); any kind of variation in coloration of tail flukes after birth year did not dramatically influence yearly sighting backgrounds. Due to the fact that whales take a trip in grslrfc.orgs and also periodically create secure social affiliations, the probcapacity of resighting individuals at any type of one time may not be independent (assumption 3 above; Weinwealthy 1991, 1998; Weinrich and also Kuhlberg 1991). However, we thought about resightings on an yearly basis, so such dependence is most likely to be reduced. Although the probcapacity of resighting and opportunity for calf mortality are connected to that of their mothers, the reality that we analyzed the 2 teams individually precludes any kind of potential problems of nonindependence. Individuals were considered to be nonadults from birth until their fifth year of life (Clapham and Mayo 1990) or until they produced a calf.

Model selection followed the step-dvery own methodology suggested by Lebreton et al. (1992), by modeling resighting probabilities first to retain as a lot power as possible for tests of survival parameters. We assessed the results of year (y) and haplotype (h) on resighting prices to test for genetically based variation and also to account for potential variation in observer effort from year to year. We likewise assessed the results of haplokind (h), clade level V (c), year (y), and age (a) on survival for female calves. Age effects on resighting and also survival were not thought about for adults bereason we had no a priori criteria for delimiting systematic age classes. The effects of year (and also age) were modeled first to preserve power for tests by maternal lineage haplotype. The model framework through the lowest Akaike Indevelopment Criterion (AIC) was selected as the most most likely model (Lebreton et al. 1992). Formal tests of certain results provided LR tests in between nested models. Model selection for calves provided modified full capture backgrounds that were coded as zero after 5 years (termed dummy variables). Subsequent analyses ignored results based upon ages better than 5 years.

Projections of Genetic Variation

Fecundity and also survivorship rates were provided to construct a basic deterministic model to project likely population trajectories for individual haplotypes within the Gulf of Maine population. The model considered females only and also was based upon postbreeding censsupplies, as the actual data were built up this means. Two age classes, calf and adult, that differed in survival prices and fecundity were provided. The version included the fecundity and also survival worths figured out for each haplotype from the analyses explained above.

The design rests on 6 assumptions: (1) males carry out not influence the survival and also reproduction of females, (2) prices of survival and fecundity are constant among people within an era course and among years, (3) breeding occurs in a single birth pulse, (4) the sex ratio of offspring is 50:50 and also does not vary among haplotypes, (5) the population is close to a stable age distribution, and (6) parameters are not thickness dependent. The second assumption in certain is unmost likely to be true, as both fecundity and also survival can differ among females and across years (check out Results). Limited proof additionally shows that maternal condition among some females may affect offspring sex ratio (Wiley and Clapham 1993). However before, slight violations of these assumptions are unmost likely to affect the qualitative outcomes of the design (Noon and Sauer 1992). Populations were projected for 75 years from the existing, start via an initial population of 100 people distributed in a stable age circulation. Numbers of people within each maternal family tree in the starting populace approximated the overall propercent of adults known for each maternal lineage taped throughout the examine period.

Results

Genealogy

Maternal lineage haplokinds were established for a total of 323 pets, either by direct sequencing (n = 159) or by assignment based on known relationships along matrilines (n = 164), including recent births and deceased people. A total of 23 variable and indevelopmental nucleotide sites identified 19 unique haplokinds within the populace of known and also sampled individuals from the Gulf of Maine (Figure 2). Haplotypes were continuous in form and number via Baker et al. (1993, 1994a) and also Palsbøll et al. (1995). Direct sequencing of a larger percentage of the mtDNA regulate area, and an extra representative sample size presented right here, reveals better haplokind diversity than observed in earlier studies (Baker and also Medrano-Gonzalez 2002) (Figure 2). Fifteen additional sites were discovered to be polymorphic via respect to a North Pacific humpback whale and/or the homologous fin whale sequence. Whales that possess the very same haploform have been grslrfc.orged together by letter and numerical desigcountry. Haploforms are labeled so they might be linked by one or more mutational measures to the following haplotype.

The strict consensus family tree of haplotypes is shown in Figure 3. Stepwise addition heuristic or branch and also bound searches of haplotypes caused 15 equally parsimonious trees, each containing 37 steps (confidence interval =.622; RI =.860). Bootstrap worths and also degeneration indices offered assistance for the 2 principal clades containing BCD and also IJK lineperiods. The topology for the mtDNA manage region family tree shown in Figure 3 is continuous via that of Baker et al. (1994a). The present and also previous researches (Baker et al. 1994a; Palsbøll et al. 1995) perform not test whether any kind of of the maternal lineeras are associated with particular geographical locations within the Gulf of Maine.

Relationships of Fecundity With Genealogy

There was no substantial difference in intend fecundity (number of calves per female audit for heterogeneity of recapture probcapability over the research period) in between the females discovered in the 2 clades (level V; Kruskal–Wallis test, χ2 = 0.028; df = 1; P =.867). The circulation of fecundity across low (0–0.25), medium (0.25–0.5), and also high (higher than 0.5) categories likewise did not differ among the two major clades (level V; Fisher"s precise test, P =.133).

We tested individual maternal lineperiods and various other clades for differential fecundity over the research duration. Based on fecundity, the propercent of people differed among the grslrfc.orgings of maternal lineperiods. There were no substantial differences discovered among level I lineages (Fisher"s specific test = 0.630) or level IV lineages within the BCD clade (Fisher"s specific test = 0.300). Huge differences in fecundity were detected among females at level II (Fisher"s specific test = 0.052), level III (0.019), and level IV lineages within the IJK clade (Fisher"s precise test = 0.048).

Examicountry of level II lineage grslrfc.orgings revealed that females with the highest fecundity (probability of being oboffered with a calf > 0.5) were existing in maternal family tree C (33%) and clade I-I1-I2 (24%). Clades D1-D, J-J1-J2-J3, and also K2-K3 had actually no females that would be identified through high refertile rates. Individual females with the lowest fecundity (probability of being observed with a calf 2 = 28.01; df = 16; P =.032). When a hereditary component is consisted of to test reproductive success by year for the two clades (BCD versus IJK), a solid and raising temporal trfinish was oboffered in fecundity in redeveloping females of the IJK clade (F = 23.9; df = 1; P =.002), flourishing steadily by a product of 1.022 every year (Figure 4). No trend was oboffered among females of the BCD clade (F = 0.372; df = 1; P =.553).

Patterns of Survival and also Resighting History

For calf survival, the version with the finest fit (as defined by the lowest AIC value) emerged wright here resighting probabilities for calves varied through age (Table 1; model 7). Tbelow was no significant effect of year by haplokind interactivity on resighting or any results linked through maternal lineage on survival. Individuals were much less likely to be viewed throughout their first postweaning year than between the periods of 2–5 years (Table 2). Calf survival varied among years, yet was not considerably influenced by age among the lineeras analyzed (Table 1). The best design gave an excellent fit to the oboffered resighting information (goodness-of-fit test, χ2 = 2.28; df = 16; P =.999). Annual survival price averaged 0.702 over 16 years, and ranged from 0.142 to 1.000 (Table 2). The lowest calf survival approximates emerged between 1993 and 1995.

The best-fitting design for adult female survival shown that resighting differed considerably among years and survival was constant (Table 3, model 7). A version incorporating annual variation in resighting prices offered a significantly much better fit to the data than otherwise identical models through haplotype-dependent resighting (i.e., encountering specific lineeras preferentially to others) or a consistent price of resighting (Table 3, models 1 and also 2 versus 3). Models including variation in survival based on year, haplokind, and clade additionally faibrought about carry out a significantly much better fit than did a less complicated version via a continuous, prevalent survival rate (Table 3, models 4, 5, and 6 versus 7). The approximated annual survival of adult female whales in the Gulf of Maine was high: 0.96. Based on the a lot of most likely model, survival and also resighting prices for adult whales did not differ substantially in between different haploforms or among years (Table 4). The finest model for adult prices provided an excellent fit to the data based upon a goodness-of-fit test (χ2 = 5.27; df = 16; P =.994).

Population Projection and Genetic Variation

The projections for Gulf of Maine humpago whales, based upon the empirically established survival, resighting, and also reabundant rates suggest that the populace will rise over the next 75 years. However, the fate of individual haploforms differed because of the differential fecundity among haplotypes (Figure 5). The B family tree disappeared completely from the populace. The C, D1, J, J1, J3, J4, K, and also K1A lineeras persisted, yet declined. In contrast, the D, I, I1, K2, and also K3 lineeras raised with time. After 75 years, the proportion of the female population making up the BCD clade decreased 11%, from 33% to 22% of the population, while the propercent of the populace consisting of the IJK clade boosted 30%.

Discussion

For the the majority of component, research studies including life background and genetic framework, specifically in cetaceans, have actually been carried out separately from one another. This study represents a in-depth testimonial of the correlation of life time reproductive success with fads of genetic variation in whales. Such associations at this particular level are commonly detected in the context of controlled breeding experiments of organisms typically with shorter generation times or among isolated human populations through thorough demographic documents (Austerlitz and also Heyer 1998; Roff 1992). These experiments regularly measure the influence of schosen quantitative traits on life history (Leroi et al. 1994a,b; Roff and Mousseau 1987). In related occupational, correlations in between heritability of refertile success among various sexes have actually additionally displayed a manifeterminal of heterogeneity in phenotypic distinctions within a population (Sinervo and also Zamudio 2001). Previous researches among whales have drawn web links between life background and also genetic diversity, but these have concentrated on the influence of migratory behavior over a whole population quite than on individual life-background traits (Baker et al. 1990; Palsbøll et al. 1995). Here we examine whether an association of fecundity arisen within natural grslrfc.orgings of maternal lineperiods throughout the ancestry. These associations might be attributed to refertile success having actually some underlying correlation through genetic or social maternal heritability, environment, or interaction between the 2 components. Given neutral concept, straight selective results linked with this mitochondrial marker are unlikely, but cannot be totally excluded. The opportunity of hereditary imprinting or maternal effects (noncultural) that produce distinctions in fecundity between the clades cannot be excluded as well. While a genetically or culturally inherited system for differential reabundant success is not interpreted, maternal lineage variation and also life-history characteristics show up to be correlated to differing levels of evolutionary and also eco-friendly scales.

Association of Fecundity With Genealogy

Variation in the mtDNA regulate area permits for the designation of people as members of a certain maternal family tree. These maternal lineeras are provided to partition variation in life-history parameters. In addition, the ancestry of mtDNA lineperiods is provided to guide the correct hierarchical levels of evaluation for trial and error higher-level correlations via life background. Based on the genetic outcomes and availcapability of demographic information, we felt that there were 5 major levels suitable for examicountry based upon hierarchical levels within the ancestry. While no considerable fads in fecundity were uncovered at the lowest and highest levels (levels I and also V), comparisons of fecundity based on resolved clades in the ancestry revealed that statistically substantial differences carry out exist between natural grslrfc.orgings of these maternal lineeras (levels II, III, and also part of IV).

At the level of the 2 strongly sustained clades (level V), a regression of fecundity by year reveals that the IJK clade mirrors a very substantial upward trfinish over time from 1979 to 1995, while the lineperiods within the BCD clade reprimary constant (Figure 4). Because fecundity of the IJK clade was low family member to the BCD clade early in the examine and high late in the research, comparisons of suppose differences and also distributions of fecundity worths were not substantially different between clades. We would, but, expect them to end up being considerable over time need to the clade-connected trends we oboffered in fecundity proceed as presented in Figure 4. These results contradict a lack of considerable findings in yat an early stage refertile success by Barlow and Clapham (1997) based on noticeable birth rate. Such trends in life-background features end up being apparent as soon as consideration is given to corconnections via genetic framework or various other appropriate variables.

Differences in reabundant success among people are a potential reason for change in gene frequencies over a short time period in humans (Austerlitz and Heyer 1998). This process appears to be similar to fecundity differences in between recreating females and their female offspring in Gulf of Maine humpago whales. Similar explacountries have actually been invoked for explaining unequal circulation of mitochondrial haplotypes in a huguy populace, as well as the boost in frequency of illness genes from reduced initial frequencies (Murray-McIntosh et al. 1998; Risch et al. 1995). Scribner et al. (1993) suggested that a correlation between reproductive success and hereditary variation in a bog turtle may reason tempdental alters in allele frequency. However before, no clear trfinish in allele frequency over time was detected. Variation in reproductive success among a finite variety of adults led to heterogeneity of maternal lineages of the Pacific oyster, which might not be attributed to geographical differences in spawning populaces (Li and Hedgecock 1998). Similarly our results, reflecting an association between reabundant success and also specific maternal lineperiods, cannot be attributed to geographical nonself-reliance. This is demonstrated by a absence of considerable findings of haplotype-dependent resighting models and also a result of genealogical structure for our dataset consistent through those from other researches of this populace (Baker et al. 1993; Palsbøll et al. 1995).

Variation in fecundity along maternal lineeras has actually prodiscovered after-effects for the frequency and distribution of maternal haplokinds within the Gulf of Maine, particularly as soon as such effects are linked with one component of the populace (Weir and Cockerham 1984). Barlow and also Clapham (1997) reported an asymptotic rate of rise of 1.065 for this populace based upon estimated birth interval, first-birth probcapacity, and also survival rates. However, any kind of rate of boost appears to be partitioned alengthy maternal lineeras, via one segment of the populace raising (IJK) due to lineages with greater fecundity and the various other continuing to be reasonably consistent (BCD) (Figure 4).

Population modeling reflects that a recirculation of maternal lineages for this populace is expected based on existing parameters and the association of life history via ancestry. After 75 years, assuming current rates and patterns, tbelow is a decrease in maternal family tree frequencies, via the BCD clade reflecting diminished frequency in the population. In addition, a number of lineages (maternal lineages primarily from the IJK clade) boost comparatively within the entire population because of the raised refertile success proficient among females for some of these matrilines. The projected change of haplotype frequencies and potential reduction in hereditary diversity in this populace is just exposed by examination of differences in refertile success among maternal lineperiods. The forecast of family tree persistence and extinction is based upon present prices and also trends detected in our analyses. The modeling is not intended to be predictive, but is supplied to demonstrate exactly how family tree vicapacity can differ over time. Factor to consider of temporal transforms in ecological conditions, stochastic results, or other spurious events could obviously change maternal family tree projections.

Assessing Population Effects Based on Fecundity/Genealogy Associations

Recent hypotheses indicate that a selective advantage associated through matrilines, possibly at one more locus in disequilibrium via certain maternal lineperiods, will dramatically minimize a population"s hereditary variation (Whitehead 1998). Whitehead invoked differential fitness among matrilines led to by matrilineally transmitted cultural traits to explain a reduction in hereditary diversity. Our outcomes directly demonstrate differential reabundant success among matrilines, and also empirically display its impacts on mtDNA lineperiods in the populations. Furthermore, the existence of high diversity in humpback whales worldwide and at the oceanic level suggests that various other mechanisms, such as mutational processes or large-scale migratory shifts because of climate adjust, should be taken into consideration to account for maintenance of diversity despite transforms predicted in this article based on variable reabundant success.

In among the critiques of Whitehead"s (1998) concept of selective heritcapability, Tiedemann and Milinkovitch (1999) have actually displayed that sperm whales exhilittle bit stochastic differences in remanufacturing via time and room, which boosts the variance in reabundant success among haplotype matrilines. The likelihood of such a phenomenon seemed plausible based upon modeling (Whitehead 1999), however variability of fecundity between matrilines had actually yet to be empirically demonstrated before the present research.

The effects of stochastic distinctions or ecological factors along martrilines of humpearlier whales in this research are an equally viable different, however are tough hypotheses to test. In the absence of extremely substantial geographical structuring of matrilines of humpearlier whales on the feeding grounds, these stochastic events can be subtle environmental shifts that affect fecundity (questioned additionally below) or various other nonspatial determinants, such as illness, reflected by substantial correlations in between fecundity and also matrilines (Whitehead 1999). Additionally, any kind of stochastic affect (spatial or nonspatial) that reasons variance in fecundity alengthy matrilines might take place alengthy the migratory course or on the wintering grounds of this species.

With the potential for tempdental or spatial heterogeneity, it is difficult to distinguish whether selection (genetics), stochasticity (environment), or an interaction in between the 2 has resulted in heterogeneity in fecundity among maternal lineeras of humpback whales. Direct genetic heritcapacity of fecundity and also its results on Ne have been suggested in various other species (Nei and Murata 1966), however is considered to be low (Charlesworth 1994; Nunney 1995; Roff and also Mousseau 1987). The outcomes of the present research imply that a 2.2% adjust in fecundity per year might account for the differences in maternal family tree haplotype frequency. Based on our outcomes over the 16-year period and also projected patterns, the predicted impacts on amount and kind of maternal family tree diversity persisting right into future generations (displayed in Figure 5) would certainly be continuous via effects presented or debated in Austerlitz and also Heyer (1998) and also Whitehead (1998).

Differences in cooperative feeding in areas of high prey, migratory habits, condition, and also ecological disturbances may have actually brought about variance in fecundity among the various genealogical levels. The valuable or detrimental effects conveyed from such incidents might be due to spatial grslrfc.orging of maternal relatives. Some create of hereditary heritability cslrfc.orgled via parallel social inheritance of possibly adaptive feeding layouts (Weinwell-off et al. 1992) and also putative habitat stratification of various age classes (Weinrich et al. 1997) could carry out reresource benefits among lineages that would certainly be reflected in reabundant success. On larger geographical scales, previous researches have argued female fidelity to feeding grounds. Differential conditions in certain locations within or between feeding grounds can have an impact on the differential refertile success of matrilines. Over a duration of two years, these possible eco-friendly determinants or stochastic procedures might account for the association of refertile differences via maternal lineages among humpearlier whales in the Gulf of Maine.

Patterns of Resighting and also Survival

There was no substantial finding of heterogeneity in survival probabilities alengthy maternal lineperiods for either adults or calves during the research period. While adult and calf survival do not appear to be affected by maternal family tree framework, the all at once patterns in resighting and survival are equivalent to those approximated based upon a model of interbirth intervals (Barlow and also Clapham 1997). Any impacts attributed to maternal lineage haplokind or clade perform not appear to affect yat an early stage resighting or survival. Potential results of haplotype or clade on within-year resighting probability are beyond the scope of the current study.

However before, substantial variation in regards to yearly on effects on sighting and also survival probabilities for calves and also yat an early stage effects on sighting probabilities for adults means that other determinants past maternal family tree affect the circulation and abundance of whales in this area. Shifts in prey abundance and also area in between 1992 and also 1995 in the Gulf of Maine led to a momentary abandonment of Stellwagen Bank by humpago whales (Weinrich et al. 1997), as shown by a decrease in adult resightings in 1992–93 with 1994–95. Calf survival likewise declined moderately in 1992–93 and precipitously for the 2 years (1993–95) following the first detectable shifts in abundance and also adult sightings. According to these models, calf survival in between 1993 and 1995 dropped to a 16-year low intend value of 0.1568. However, it is tough to discriminate in between this hypothesis and also preferential habitat use of Jeffrey"s Ledge by adults fairly than juveniles, for this reason confounding calf survival with sampling predisposition of calves (Weinwealthy 1998). In addition, lower probabilities for photographic identification of calves can affect interpretation of calf survival. One possible biological explacountry is that prey form and distribution may have actually more of a far-reaching impact on calf survival and adult resighting than simply leading to a short-lived relocation to another feeding area. The incapacity of adult females to fulfill power requirements in this region can have led to the decrease of resighting probability and also maybe had actually also higher after-effects for providing adequate nutrition for offspring.

Environpsychological disturbances, such as documented shifts in prey abundance, may have actually likewise effected various other patterns in survival and resighting. The mortality occasion that emerged in 1987, in enhancement to having a nonrandom distribution among maternal lineeras (Baker et al. 1994b), preyielded the lowest apparent pregnancy price for females the complying with year (Barlow and Clapham 1997). These kinds of environmental shifts might appear to take place even more typically than Barlow and also Clapham (1997) indicate and have the right to have complicated implications for resighting prices and calf survival. Such results are only identifiable once age course and also year are taken into consideration as independent components of humpback whale survival.

Our outcomes show that genetic variation can be associated through environmental shifts, resighting prices, yearly survival, and reabundant success. Crude transforms in survival regularly lead to a search for underlying causative results. Use of added parameters in estimation models past yearly variation or age course might aid rise accuracy and also reduce sources of uncertainty in survival approximates. More structured models that encompass a hereditary component have the right to even more illuminate trends in survival and also growth rate, specifically in species whose social structure and company are very correlated with family tree or phylogeny.

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The technique defined here can be used to account for distinctions in reproductive success among a varied team of long-lived species (Clutton-Brock 1988; Cooke and Rockwell 1988; Hamilton et al. 1998; LeBoeuf and Reiter 1988; Morin et al. 1994; Packer et al. 1988). The results have ramifications for conservation initiatives by demonstrating that subtle differences in life-background traits among people deserve to alter specific facets of population genetic viability. The heterogeneity of life-background traits associated through populace framework (i.e., maternal lineages) suggests that detecting patterns in survival and also resighting, and also evaluating minimum and also effective population sizes, requires detailed analyses. In many kind of instances where trends in survivorship, fecundity, and resighting are experienced, the demographic models provided to evaluate such trends have not taken into consideration the impacts of hereditary framework (i.e., Caswell et al. 1999). As demonstrated by our results, the testimonial of life-history parameters in the context of relevant genetic data uses a unique means to evaluate grslrfc.orgs or cohorts of pets that share certain traits or might be exposed to various levels and also types of stochastic impacts. The capability to detect substantial patterns by evaluating and also incorporating genetic and demographic information deserve to administer enlightening information for conservation and administration objectives. This is specifically crucial in evaluating the source of variation for oboffered and also detected fads in the lack of evident answers.