Papers by Richard Frankham

Biological Conservation, Jan 1, 2006
The role of inbreeding depression in the extinction of wild populations is controversial, largely... more The role of inbreeding depression in the extinction of wild populations is controversial, largely because there are no quantitative estimates of its impact using realistic levels of inbreeding depression. To address this deficiency, this study (1) provides a comprehensive estimate of the impact of inbreeding depression on wild, mammalian and avian species via a meta-analysis, and (2) determines the impact of this level of inbreeding depression on extinction risk over a broad taxonomic range via stochastic computer projections with and without inbreeding depression for populations with carrying capacities of 100, 500 and 2000 individuals. An average overall effect of 12 diploid lethal equivalents was found across the life-history of the species in the meta-analysis. In the stochastic computer projections, 12 diploid lethal equivalents of inbreeding depression (with purging) decreased median times to extinction by an average of 37%. These decreases were significant and of very similar magnitude, regardless of the carrying capacity modelled. Disregarding the influence of inbreeding depression on extinction risk will lead to serious overestimates of the survival prospects of threatened mammalian and avian taxa. Further, inappropriate recovery plans may be instituted if the causes of extinction risk and their relative contributions are not recognized.
Trends in Ecology & Evolution, 2013
Biological Conservation, 2014
Conservation Genetics, 2002
Human activities are simultaneously decreasing the size of wildlife populations (causing inbreedi... more Human activities are simultaneously decreasing the size of wildlife populations (causing inbreeding) and increasing the level of stress that wildlife populations must face. Inbreeding reduces population fitness and increases extinction risk. However, very little information on the impact of stressful environments on extinction risk under inbreeding is available. We evaluated the impact of full sib inbreeding on extinction risk, using
Direct estimates of extinction risk are frequently unavailable, so surrogate indicators are used ... more Direct estimates of extinction risk are frequently unavailable, so surrogate indicators are used in threatened species categori- zations, but there is inadequate information on which best predict vulnerability. We compared the ability of 16 frequently-used factors to predict extinction risk for 45 vertebrate taxa. Median times to extinction were estimated using taxon-specific stochastic population models. Population size (N) and trend
Conservation Genetics, 2001
Deleterious alleles may be removed (purged) bynatural selection in populations undergoinginbreedi... more Deleterious alleles may be removed (purged) bynatural selection in populations undergoinginbreeding. However, there is controversyregarding the effectiveness of purging inreducing the extinction risk due to inbreeding,particularly in conservation contexts. Weevaluated the effects of purging on theextinction risk due to inbreeding in Drosophila melanogaster using two basepopulations, an outbred population (non-purged)and four-way crosses between highly inbredlines derived from the same population(purged).
Introduction to Conservation Genetics written by R. Frankham, J.D. Ballou, and D.A. Briscoe, is a... more Introduction to Conservation Genetics written by R. Frankham, J.D. Ballou, and D.A. Briscoe, is a comprehensive introductory text that provides an enlightening synthesis of data and theory from ecology, population genetics, evolution, and conservation biology. The book focuses on the science of conservation genetics and is appealing in its general lack of overt advocacy, while the final section provides meaningful applications to practical problems in conservation. Important concepts outlined in the text are emphasized by case-studies taken from primary literature, and chapters are concluded with a summary followed by exercises or problems. Beginning and seasoned researchers alike will benefit from a reference copy on their shelves.
Conservation Biology, 1992
... Sydney, NSW 2109, Australia JM MALPICA* A. ROBERTSONt Institute of Animal Genetics West Mains... more ... Sydney, NSW 2109, Australia JM MALPICA* A. ROBERTSONt Institute of Animal Genetics West Mains Road Edinburgh EH93JN, scotland, UI~ GABRIELLE J. SMITH ... NerN has been reported to be about 0.4-0.5 in humans (Emigh & Pollak 1979; Wood 1987), 0.75 in the snail ...

Zoo Biology, 1992
Immigration into small isolated captive and wild populations is recommended to alleviate inbreedi... more Immigration into small isolated captive and wild populations is recommended to alleviate inbreeding depression. The effects on reproductive fitness of introducing one immigrant into 10 small partially inbred captive populations of D. rnelunogaster were evaluated. The relative reproductive fitness of the immigrant populations (0.628) was approximately double that of the isolated populations (0.294) and about halfway between the isolated populations and the outbred base population ( 1 .OO). Every replicate population increased in fitness following the introduction of an immigrant. The improvements in reproductive fitness shown by the immigrant populations were not due to F, hybrid vigor, as the experimental populations underwent three generations of random mating prior to the fitness tests. These results indicate substantial benefits can be gained by the translocation of as few as a single animal between small, partially inbred populations. 0 1992 Wiley-Liss, Inc.
Zoo Biology, 1997
Captive populations of endangered species are managed to preserve genetic diversity and retain re... more Captive populations of endangered species are managed to preserve genetic diversity and retain reproductive fitness. Minimizing kinship (MK) has been predicted to maximize the retention of gene diversity in pedigreed populations with unequal founder representation. MK was ...

Proceedings of the National Academy of Sciences, 2004
There is controversy concerning the role of genetic factors in species extinctions. Many authors ... more There is controversy concerning the role of genetic factors in species extinctions. Many authors have asserted that species are usually driven to extinction before genetic factors have time to impact them, but few studies have seriously addressed this issue. If this assertion is true, there will be little difference in genetic diversity between threatened and taxonomically related nonthreatened species. We compared average heterozygosities in 170 threatened taxa with those in taxonomically related nonthreatened taxa in a comprehensive metaanalysis. Heterozygosity was lower in threatened taxa in 77% of comparisons, a highly significant departure from the predictions of the no genetic impact hypothesis. Heterozygosity was on average 35% lower (median 40%) in threatened taxa than in related nonthreatened ones. These differences in heterozygosity indicate lowered evolutionary potential, compromised reproductive fitness, and elevated extinction risk in the wild. Independent evidence from stochastic computer projections has demonstrated that inbreeding depression elevates extinction risk for threatened species in natural habitats when all other threatening processes are included in the models. Thus, most taxa are not driven to extinction before genetic factors affect them adversely.

PLoS ONE, 2010
The persistence of tropical coral reefs is threatened by rapidly increasing climate warming, caus... more The persistence of tropical coral reefs is threatened by rapidly increasing climate warming, causing a functional breakdown of the obligate symbiosis between corals and their algal photosymbionts (Symbiodinium) through a process known as coral bleaching. Yet the potential of the coral-algal symbiosis to genetically adapt in an evolutionary sense to warming oceans is unknown. Using a quantitative genetics approach, we estimated the proportion of the variance in thermal tolerance traits that has a genetic basis (i.e. heritability) as a proxy for their adaptive potential in the widespread Indo-Pacific reef-building coral Acropora millepora. We chose two physiologically different populations that associate respectively with one thermotolerant (Symbiodinium clade D) and one less tolerant symbiont type (Symbiodinium C2). In both symbiont types, pulse amplitude modulated (PAM) fluorometry and high performance liquid chromatography (HPLC) analysis revealed significant heritabilities for traits related to both photosynthesis and photoprotective pigment profile. However, quantitative real-time polymerase chain reaction (qRT-PCR) assays showed a lack of heritability in both coral host populations for their own expression of fundamental stress genes. Coral colony growth, contributed to by both symbiotic partners, displayed heritability. High heritabilities for functional key traits of algal symbionts, along with their short clonal generation time and high population sizes allow for their rapid thermal adaptation. However, the low overall heritability of coral host traits, along with the corals' long generation time, raise concern about the timely adaptation of the coral-algal symbiosis in the face of continued rapid climate warming.

Genetical Research, 1996
Highly variable DNA polymorphisms called microsatellites are rapidly becoming the marker of choic... more Highly variable DNA polymorphisms called microsatellites are rapidly becoming the marker of choice in population genetic studies. Until now, microsatellites have not been utilized for Drosophila studies. We have identified eight polymorphic microsatellite loci in Drosophila melanogaster and used them to characterize the genetic variation in a wild population from the Tyrrell's winery in Australia. Microsatellites were isolated from a partial genomic DNA library. All microsatellites consist of (AC)n repeats ranging from n = 2 to n = 24. Six loci were assigned to chromosomal location by genetic mapping, with three loci on chromosome II, one locus on chromosome III and two loci on the X chromosome. Up to four microsatellite loci were multiplexed in the same reaction. Microsatellite variation is substantially greater than allozyme variation in the Tyrrell's Drosophila population. 80% of the microsatellite loci examined are polymorphic, compared with 28% of allozymes. The mean number of alleles per polymorphic locus is 5.2 in microsatellites compared with 3.0 in allozymes. The average observed heterozygosity of polymorphic microsatellites is 47% compared with 26% for allozymes. Microsatellite variation in Drosophila melanogaster is similar to that reported for other insects. Higher variability commends microsatellites over allozymes for genetic studies in Drosophila melanogaster.

Genetical Research, 2005
Quantitative genetic variation, the main determinant of the ability to evolve, is expected to be ... more Quantitative genetic variation, the main determinant of the ability to evolve, is expected to be lost in small populations, but there are limited data on the effect, and controversy as to whether it is similar to that for near neutral molecular variation. Genetic variation for abdominal and sternopleural bristle numbers and allozyme heterozygosity were estimated in 23 populations of Drosophila melanogaster maintained at effective population sizes of 25, 50, 100, 250 or 500 for 50 generations, as well as in 19 highly inbred populations and the wild outbred base population. Highly significant negative regressions of proportion of initial genetic variation retained on inbreeding due to finite population size were observed for both quantitative characters (b = -0.67 +/- 0.14 and -0.58 +/- 0.11) and allozyme heterozygosity (b = -0.79 +/- 0.10), and the regression coefficients did not differ significantly. Thus, quantitative genetic variation is being lost at a similar rate to molecular genetic variation. However, genetic variation for all traits was lost at rates significantly slower than predicted by neutral theory, most likely due to associative overdominance. Positive, but relatively low correlations were found among the different measures of genetic variation, but their low magnitudes were attributed to large sampling errors, rather than differences in the underlying processes of loss.

Evolution, 2003
The ability of populations to undergo adaptive evolution depends on the presence of genetic varia... more The ability of populations to undergo adaptive evolution depends on the presence of genetic variation for ecologically important traits. The maintenance of genetic variation may be influenced by many variables, particularly long-term effective population size and the strength and form of selection. The roles of these factors are controversial and there is very little information on their impacts for quantitative characters. The aims of this study were to determine the impacts of population size and variable versus constant prior environmental conditions on fitness and the magnitude of response to selection. Outbred and inbred populations of Drosophila melanogaster were maintained under benign, constant stressful, and variably stressful conditions for seven generations, and then forced to adapt to a novel stress for seven generations. Fitness and adaptability were assayed in each replicate population. Our findings are that: (1) populations inbred in a variable environment were more adaptable than those inbred in a constant environment; (2) populations adapted to a prior stressful environment had greater fitness when reared in a novel stress than those less adapted to stress; (3) inbred populations had lower fitness and were less adaptable than the outbred population they were derived from; and (4) strong lineage effects were detectable across environments in the inbred populations.

Evolution, 2001
The ability of populations to undergo adaptive evolution depends on the presence of quantitative ... more The ability of populations to undergo adaptive evolution depends on the presence of quantitative genetic variation for ecologically important traits. Although molecular measures are widely used as surrogates for quantitative genetic variation, there is controversy about the strength of the relationship between the two. To resolve this issue, we carried out a meta-analysis based on 71 datasets. The mean correlation between molecular and quantitative measures of genetic variation was weak (r ϭ 0.217). Furthermore, there was no significant relationship between the two measures for life-history traits (r ϭ Ϫ0.11) or for the quantitative measure generally considered as the best indicator of adaptive potential, heritability (r ϭ Ϫ0.08). Consequently, molecular measures of genetic diversity have only a very limited ability to predict quantitative genetic variability. When information about a population's short-term evolutionary potential or estimates of local adaptation and population divergence are required, quantitative genetic variation should be measured directly.

Conservation Genetics, 2000
Inbreeding and loss of genetic diversity are predicted to decrease the resistance of species to d... more Inbreeding and loss of genetic diversity are predicted to decrease the resistance of species to disease. However, this issue is controversial and there is limited rigorous scientific evidence available. To test whether inbreeding and loss of genetic diversity affect a host's resistance to disease, Drosophila melanogaster populations with different levels of inbreeding and genetic diversity were exposed separately to (a) thuringiensin, an insecticidal toxin produced by some strains of Bacillus thuringiensis, and (b) live Serratia marcescens bacteria. Inbreeding and loss of genetic diversity significantly reduced resistance of D. melanogaster to both the thuringiensin toxin and live Serratia marcescens. For both, the best fitting relationships between resistance and inbreeding were curvilinear. As expected, there was wide variation among replicate inbred populations in disease resistance. Lowered resistances to both the toxin and the pathogen in inbred populations were due to specific resistance alleles, rather than generalized inbreeding effects, as correlations between resistance and population fitness were low or negative. Wildlife managers should strive to minimise inbreeding and loss of genetic diversity within threatened populations and to minimise exposure of inbred populations to disease.
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Papers by Richard Frankham