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Jonathan Q. Richmond

Publications and source records attributed to Jonathan Q. Richmond.

34 records · Page 2Linked to original sources

Loss of dendritic connectivity in southern California's urban riverscape facilitates decline of an endemic freshwater fish

Life history adaptations and spatial configuration of metapopulation networks allow certain species to persist in extreme fluctuating environments, yet long-term stability within these systems relies on the maintenance of linkage habitat. Degradation of such linkages in urban riverscapes can disrupt this dynamic in aquatic species, leading to increased extinction debt in local populations experiencing environment-related demographic flux. We used microsatellites and mtDNA to examine the effects of collapsed network structure in the endemic Santa Ana sucker Catostomus santaanae of southern California, a threatened species affected by natural flood-drought cycles, ‘boom-and-bust’ demography, hybridization, and presumed artificial transplantation. Our results show a predominance of drift-mediated processes in shaping population structure, and that reverse mechanisms for counterbalancing the genetic effects of these phenomena have dissipated with the collapse of dendritic connectivity. We use approximate Bayesian models to support two cases of artificial transplantation, and provide evidence that one of the invaded systems better represents the historic processes that maintained genetic variation within watersheds than any remaining drainages where C. santaanae is considered native. We further show that a stable dry gap in the northern range is preventing genetic dilution of pure C. santaanae persisting upstream of a hybrid assemblage involving a non-native sucker, and that local accumulation of genetic variation in the same drainage is influenced by position within the network. This work has important implications for declining species that have historically relied on dendritic metapopulation networks to maintain source-sink dynamics in phasic environments, but no longer possess this capacity in urban-converted landscapes.

California

An extirpated lineage of a threatened frog species resurfaces in southern California

Southern California has experienced widespread amphibian declines since the 1960s. One species, the Vulnerable California red-legged frog Rana draytonii , is now considered to be extirpated from most of southern California. In February 2017 a population of R. draytonii was discovered in the southern foothills of the San Bernardino Mountains of Riverside County, California, near the edge of the species’ historical distribution. This population belongs to an mtDNA lineage that was presumed to be extirpated within the USA but is still extant in Baja California, Mexico. This discovery increases the potential for future, evolutionarily informed translocations within the southern portion of this species’ range in California.

Oryx

At the end of the line: independent overwater colonizations of the Solomon Islands by a hyperdiverse trans-Wallacean lizard lineage (Cyrtodactylus: Gekkota: Squamata)

The islands of East Melanesia have generated key insights into speciation processes and community assembly. However, when and how these islands began to form, emerge and accumulate endemic taxa remains poorly understood. Here, we show that two divergent lineages within the world’s most diverse genus of geckos ( Cyrtodactylus ) occur in the Solomon Islands. One large-bodied species is nested within a radiation from far eastern New Guinea, with inferred colonization, spread and diversification since the late Miocene. In contrast, a newly sampled and relatively small species with a restricted distribution on Guadalcanal Island is a relict that diverged from extant congeners around the early to mid-Miocene. Similar Miocene divergences from extralimital relatives have been inferred for other endemic bird, bat and lizard lineages in East Melanesia. In contrast, across all lineages (including divergent relictual lineages), there is little evidence for endemic in situ diversification within East Melanesia predating the Pliocene (especially in the Solomon Islands). While some East Melanesian endemic lineages may have origins on progenitor islands during the Miocene or even earlier, current evidence suggests the in situ diversification and assembly of extant biological communities commenced around the end of the Miocene.

Zoological Journal of the Linnean Society

Persistence of historical population structure in an endangered species despite near-complete biome conversion in California's San Joaquin Desert

Genomic responses to habitat conversion can be rapid, providing wildlife managers with time-limited opportunities to enact recovery efforts that use population connectivity information that reflects predisturbance landscapes. Despite near-complete biome conversion, such opportunities may still exist for the endemic fauna and flora of California's San Joaquin Desert, but comprehensive genetic data sets are lacking for nearly all species in the region. To fill this knowledge gap, we studied the rangewide population structure of the endangered blunt-nosed leopard lizard Gambelia sila , a San Joaquin Desert endemic, using restriction site-associated DNA (RAD), microsatellite and mtDNA data to test whether admixture patterns and estimates of effective migration surfaces (EEMS) can identify land areas with high population connectivity prior to the conversion of native xeric habitats. Clustering and phylogenetic analyses indicate a recent shared history between numerous isolated populations and EEMS reveals latent signals of corridors and barriers to gene flow over areas now replaced by agriculture and urbanization. Conflicting histories between the mtDNA and nuclear genomes are consistent with hybridization with the sister species G. wislizenii , raising important questions about where legal protection should end at the southern range limit of G. sila . Comparative analysis of different data sets also adds to a growing list of advantages in using RAD loci for genetic studies of rare species. We demonstrate how the results of this work can serve as an evolutionary guidance tool for managing endemic, arid-adapted taxa in one of the world's most compromised landscapes.

California

Historical habitat barriers prevent ring-like genetic continuity throughout the distribution of threatened Alameda Striped Racers ( Coluber lateralis euryxanthus )

We used microsatellites and mtDNA sequences to examine the mixed effects of geophysical, habitat, and contemporary urban barriers on the genetics of threatened Alameda Striped Racers (Coluber lateralis euryxanthus), a species with close ties to declining coastal scrub and chaparral habitat in the eastern San Francisco Bay area of California. We used cluster assignments to characterize population genetic structuring with respect to land management units and approximate Bayesian analysis to rank the ability of five alternative evolutionary hypotheses to explain the inferred structure. Then, we estimated rates of contemporary and historical migration among the major clusters and measured the fit of different historical migration models to better understand the formation of the current population structure. Our results reveal a ring-like pattern of historical connectivity around the Tri-Valley area of the East Bay (i.e., San Ramon, Amador, and Livermore valleys), with clusters largely corresponding to different management units. We found no evidence of continuous gene flow throughout the ring, however, and that the main gap in continuity is centered across the Livermore Valley. Historical migration models support higher rates of gene flow away from the terminal ends of the ring on the north and south sides of the Valley, compared with rates into those areas from western sites that border the interior San Francisco Bay. We attribute the break in ring-like connectivity to the presence of unsuitable habitat within the Livermore Valley that has been reinforced by 20th century urbanization, and the asymmetry in gene flow rates to spatial constraints on movement and east–west environmental gradients influenced by the proximity of the San Francisco Bay.

California

Rare alluvial sands of El Monte Valley, California (San Diego County), support high herpetofaunal species richness and diversity, despite severe habitat disturbance

We characterized the species richness, diversity, and distribution of amphibians and reptiles inhabiting El Monte Valley, a heavily disturbed, alluvium-filled basin within the lower San Diego River in Lakeside, California. This rare habitat type in coastal southern California is designated as a critical sand resource by the state of California and is currently under consideration for a large-scale sand mining operation with subsequent habitat restoration. We conducted field surveys from June 2015 to May 2016 using drift fence lines with funnel traps, coverboard arrays, walking transects, and road driving. We recorded 1,208 total captures, revealing high species richness and diversity, but with marked unevenness in species' abundances. Snakes were the most species-rich taxonomic group (13 species representing 11 genera), followed by lizards (11 species representing 9 genera). After the southern Pacific rattlesnake ( Crotalus oreganus helleri ), the California glossy snake ( Arizona elegans occidentalis ) was the second most frequently detected snake species ( n = 23 captures). Amphibian species richness was limited to only three species in three genera. Despite the relatively limited 12-month sampling period, a longstanding drought, and severe habitat disturbance, our study demonstrates that El Monte Valley harbors a rich herpetofauna that includes many sensitive species.

California

Status of the California Red-legged Frog (Rana draytonii) in the State of Baja California, México

The California Red-legged Frog ( Rana draytonii ) is a threatened species in the United States that has undergone population declines, especially in southern California. Due to the lack of information on the status of Mexican populations, we surveyed for the presence of R. draytonii in Baja California and assessed possible threats to population persistence. Our study area extended from the U.S.-Mexican border to the southern end of the distribution of the species in the Sierra San Pedro Mártir. We found R. draytonii at six of 15 historical sites, none at five proxy sites (i.e., alternative sites chosen because the historical record lacked precise locality data), and four at 24 additional sites. The 10 occupied sites are within three watersheds in the Sierra San Pedro Mártir (two sites at Arroyo San Rafael, two sites at Arroyo San Telmo, and six sites at Arroyo Santo Domingo). We did not detect R. draytonii at 60% of historical sites, including the highest elevation site at La Encantada and multiple low-elevation coastal drainages, suggesting the species has declined in Baja California. The threats we noted most frequently were presence of exotic aquatic animal species, water diversion, and cattle grazing. Management of remaining populations and local education is needed to prevent further declines.

Baja California

Invaded invaders: Infection of invasive Brown Treesnakes on Guam by an exotic larval cestode with a life cycle comprised of non-native hosts

Background Multiple host introductions to the same non-native environment have the potential to complete life cycles of parasites incidentally transported with them. Our goal was to identify a recently detected parasitic flatworm in the invasive Brown Treesnake ( Boiga irregularis ) on the remote Pacific island of Guam. We considered possible factors influencing parasite transmission, and tested for correlations between infection status and potential indicators of host fitness. We used genetic data from the parasite and information about the native ranges of other possible non-native hosts to hypothesize how it arrived on Guam and how its life cycle may be currently supported. Methods We identified the parasite by comparing larval morphology and mtDNA sequences with other Pseudophyllid tapeworms. We assessed probability of infection in individual snakes using logistic regression and examined different factors influencing presence of parasites in hosts. Results We identified the parasite as the pseudophyllid cestode Spirometra erinaceieuropaei , with all sampled worms from multiple snakes sharing a single mtDNA haplotype. Infection appears to be limited to the only freshwater watershed on the island, where infection prevalence was high (77.5%). Larger snakes had a higher probability of being infected, consistent with the chronic nature of such infections. While infection status was positively correlated with body condition, infected snakes tended to have lower intra-peritoneal fat body mass, potentially indicating a negative effect on energy stores. Conclusions We discovered that B . irregularis inhabiting a small area of forested habitat in a freshwater watershed on Guam are often infected by a novel parasite of Asian origin. While further work is needed, this species of Spirometra , itself a non-native species, likely depends on a suite of recently introduced hosts from different parts of the world to complete the life cycle. This baseline study provides little evidence of any effects on host fitness, but additional data are needed to more thoroughly explore the consequences of infection in this invasive snake population.

PLoS ONE

Niche divergence builds the case for ecological speciation in skinks of the Plestiodon skiltonianus species complex

Adaptation to different thermal environments has the potential to cause evolutionary changes that are sufficient to drive ecological speciation. Here, we examine whether climate-based niche divergence in lizards of the Plestiodon skiltonianus species complex is consistent with the outcomes of such a process. Previous work on this group shows that a mechanical sexual barrier has evolved between species that differ mainly in body size and that the barrier may be a by-product of selection for increased body size in lineages that have invaded xeric environments; however, baseline information on niche divergence among members of the group is lacking. We quantified the climatic niche using mechanistic physiological and correlative niche models and then estimated niche differences among species using ordination techniques and tests of niche overlap and equivalency. Our results show that the thermal niches of size-divergent, reproductively isolated morphospecies are significantly differentiated and that precipitation may have been as important as temperature in causing increased shifts in body size in xeric habitats. While these findings alone do not demonstrate thermal adaptation or identify the cause of speciation, their integration with earlier genetic and behavioral studies provides a useful test of phenotype–environment associations that further support the case for ecological speciation in these lizards.

Ecology and Evolution

Ephemeral stream reaches preserve the evolutionary and distributional history of threespine stickleback in the Santa Clara and Ventura River watersheds of southern California

Much remains to be understood about the evolutionary history and contemporary landscape genetics of unarmored threespine stickleback in southern California, where populations collectively referred to as Gasterosteus aculeatus williamsoni have severely declined over the past 70+ years and are now endangered. We used mitochondrial sequence and microsatellite data to assess the population genetics and phylogeography of unarmored populations sampled immediately downstream from the type locality of G. a. williamsoni in the upper Santa Clara River, and assessed their distinctiveness with respect to low-armor populations in the downstream sections of the river and the adjacent Ventura River. We also characterized the geographic limits of different plate morphs and evaluated the congruence of those boundaries with barriers to dispersal in both river systems and to neutral genetic variation. We show substantial population structuring within the upper reach of the Santa Clara River, but little partitioning between the lower Santa Clara and Ventura Rivers—we attribute these patterns to different ancestry between spatially subdivided populations within the same drainage, a predominance of downstream gene flow, and ability for coastal dispersal between the Santa Clara and Ventura Rivers. We also show that alleles from introduced low-plate stock have infiltrated a native population in at least one upper Santa Clara River tributary, causing this formerly unarmored population to become gradually low-plated over a 30 + year time period. Measures of genetic diversity, census surveys, and severe habitat disturbance all indicate that unarmored stickleback near the type locality are currently at high risk of extinction.

California

Testing for multiple invasion routes and source populations for the invasive brown treesnake ( Boiga irregularis ) on Guam: implications for pest management

The brown treesnake ( Boiga irregularis ) population on the Pacific island of Guam has reached iconic status as one of the most destructive invasive species of modern times, yet no published works have used genetic data to identify a source population. We used DNA sequence data from multiple genetic markers and coalescent-based phylogenetic methods to place the Guam population within the broader phylogeographic context of B. irregularis across its native range and tested whether patterns of genetic variation on the island are consistent with one or multiple introductions from different source populations. We also modeled a series of demographic scenarios that differed in the effective size and duration of a population bottleneck immediately following the invasion on Guam, and measured the fit of these simulations to the observed data using approximate Bayesian computation. Our results exclude the possibility of serial introductions from different source populations, and instead verify a single origin from the Admiralty Archipelago off the north coast of Papua New Guinea. This finding is consistent with the hypothesis that B. irregularis was accidentally transported to Guam during military relocation efforts at the end of World War II. Demographic model comparisons suggest that multiple snakes were transported to Guam from the source locality, but that fewer than 10 individuals could be responsible for establishing the population. Our results also provide evidence that low genetic diversity stemming from the founder event has not been a hindrance to the ecological success of B. irregularis on Guam, and at the same time offers a unique ‘genetic opening’ to manage snake density using classical biological approaches.

Biological Invasions

Population declines lead to replicate patterns of internal range structure at the tips of the distribution of the California red-legged frog ( Rana draytonii )

Demographic declines and increased isolation of peripheral populations of the threatened California red-legged frog ( Rana draytonii ) have led to the formation of internal range boundaries at opposite ends of the species’ distribution. While the population genetics of the southern internal boundary has been studied in some detail, similar information is lacking for the northern part of the range. In this study, we used microsatellite and mtDNA data to examine the genetic structuring and diversity of some of the last remaining R. draytonii populations in the northern Sierra Nevada, which collectively form the northern external range boundary. We compared these data to coastal populations in the San Francisco Bay Area, where the species is notably more abundant and still exists throughout much of its historic range. We show that ‘external’ Sierra Nevada populations have lower genetic diversity and are more differentiated from one another than their ‘internal’ Bay Area counterparts. This same pattern was mirrored across the distribution in California, where Sierra Nevada and Bay Area populations had lower allelic variability compared to those previously studied in coastal southern California. This genetic signature of northward range expansion was mirrored in the phylogeography of mtDNA haplotypes; northern Sierra Nevada haplotypes showed greater similarity to haplotypes from the south Coast Ranges than to the more geographically proximate populations in the Bay Area. These data cast new light on the geographic origins of Sierra Nevada R. draytonii populations and highlight the importance of distinguishing the genetic effects of contemporary demographic declines from underlying signatures of historic range expansion when addressing the most immediate threats to population persistence. Because there is no evidence of contemporary gene flow between any of the Sierra Nevada R. draytonii populations, we suggest that management activities should focus on maintaining and creating additional ponds to support breeding within typical dispersal distances of occupied habitat.

Biological Conservation

Evolutionary dynamics of a rapidly receding southern range boundary in the threatened California red-legged frog ( Rana draytonii )

Populations forming the edge of a species range are often imperiled by isolation and low genetic diversity, with proximity to human population centers being a major determinant of edge stability in modern landscapes. Since the 1960s, the California red-legged frog ( Rana draytonii ) has undergone extensive declines in heavily urbanized southern California, where the range edge has rapidly contracted northward while shifting its cardinal orientation to an east-west trending axis. We studied the genetic structure and diversity of these frontline populations, tested for signatures of contemporary disturbance, specifically fire, and attempted to disentangle these signals from demographic events extending deeper into the past. Consistent with the genetic expectations of the ‘abundant-center’ model, we found that diversity, admixture, and opportunity for random mating increases in populations sampled successively further away from the range boundary. Demographic simulations indicate that bottlenecks in peripheral isolates are associated with processes extending tens to a few hundred generations in the past, despite the demographic collapse of some due to recent fire-flood events. While the effects of recent disturbance have left little genetic imprint on these populations, they likely contribute to an extinction debt that will lead to continued range contraction unless management intervenes to stall or reverse the process.

California

Selection, trans-species polymorphism, and locus identification of major histocompatibility complex class IIβ alleles of New World ranid frogs

Genes encoded by the major histocompatibility complex (MHC) play key roles in the vertebrate immune system. However, our understanding of the evolutionary processes and underlying genetic mechanisms shaping these genes is limited in many taxa, including amphibians, a group currently impacted by emerging infectious diseases. To further elucidate the evolution of the MHC in frogs (anurans) and develop tools for population genetics, we surveyed allelic diversity of the MHC class II ??1 domain in both genomic and complementary DNA of seven New World species in the genus Rana (Lithobates). To assign locus affiliation to our alleles, we used a "gene walking" technique to obtain intron 2 sequences that flanked MHC class II?? exon 2. Two distinct intron sequences were recovered, suggesting the presence of at least two class II?? loci in Rana. We designed a primer pair that successfully amplified an orthologous locus from all seven Rana species. In total, we recovered 13 alleles and documented trans-species polymorphism for four of the alleles. We also found quantitative evidence of selection acting on amino acid residues that are putatively involved in peptide binding and structural stability of the ??1 domain of anurans. Our results indicated that primer mismatch can result in polymerase chain reaction (PCR) bias, which influences the number of alleles that are recovered. Using a single locus may minimize PCR bias caused by primer mismatch, and the gene walking technique was an effective approach for generating single-copy orthologous markers necessary for future studies of MHC allelic variation in natural amphibian populations. ?? 2010 Springer-Verlag.

Immunogenetics

Delayed genetic effects of habitat fragmentation on the ecologically specialized Florida sand skink ( Plestiodon reynoldsi )

Populations rarely show immediate genetic responses to habitat fragmentation, even in taxa that possess suites of traits known to increase their vulnerability to extinction. Thus conservation geneticists must consider the time scale over which contemporary evolutionary processes operate to accurately portray the effects of habitat isolation. Here, we examine the genetic impacts of fragmentation on the Florida sand skink Plestiodon reynoldsi , a sand swimming lizard that is highly adapted to the upland scrub habitat of central Florida. We studied fragments located on the southern Lake Wales Ridge, where human activity in the latter half of the 20th century has modified the natural patchiness of the landscape. Based on a relaxed molecular clock method, we estimate that sand skinks have persisted in this region for approximately 1.5 million years and that the time frame of human disturbance is equivalent to fewer than 30 skink generations. Using genotypes from eight microsatellite loci, we screened for molecular signatures of this disturbance by assessing congruence between population structure, as inferred from spatially-informed Bayesian assignment tests, and the current geography of scrub fragments. We also tested for potential intrapopulation genetic effects of inbreeding in isolated populations by comparing the average pairwise relatedness of individuals within fragments of different areas and isolation. Our results indicate that although some patches show a higher degree of relatedness than expected under random mating, the genetic effects of recent isolation are not evident in this part of the species’ range. We argue that this result is an artefact of a time-lag in the response to disturbance, and that species-typical demographic features may explain the genetic inertia observed in these populations.

Conservation Genetics