Coqui frog invasions change invertebrate communities in Hawaii

The Puerto Rican coqui frog (Eleutherodactylus coqui) invaded Hawaii in the late 1980s. Because the coqui reaches high densities and consumes large quantities of invertebrates, it was hypothesized to change invertebrate communities where it invades. Previous research found that coquis can change invertebrate communities, but these studies used highly manipulative, small-scale experiments. The objective of this research was to determine whether coquis create community-level changes in invertebrate communities at the landscape scale. We collected leaf litter, flying, and foliage invertebrates on both sides of 15 coqui invasion fronts across the island of Hawaii. Multivariate analyses show that coquis are associated with changes in leaf-litter communities, primarily reductions in Acari, but are not associated with overall changes in flying or foliage communities. Across sites, coquis reduced the total number of leaf-litter invertebrates by 27%, specifically by reducing Acari by 36%. Across sites, coquis increased flying Diptera by 19%. Changes were greater where coqui densities were higher. We suggest that coquis changed leaf-litter communities primarily through direct predation, but that they increased Diptera through the addition of frog carcasses and excrement. Results support previous studies conducted in more controlled settings, but add to our understanding of the invasion by showing that coqui effects on invertebrate communities are measurable at the landscape scale.     

Potential consequences of the coqui frog invasion in Hawaii

The Puerto Rican frog, Eleutherodactylus coqui, has invaded Hawaii and has negatively impacted the state's multimillion dollar floriculture, nursery and tourist industries; however, little is known about the ecological consequences of the invasion. Using data from Puerto Rico and Hawaii, the authors summarize the potential consequences of the invasion and describe future research needs. It could be predicted that the coqui would reduce the abundance of Hawaii's endemic invertebrates. However, data suggest that coquis are mostly consuming non‐native invertebrates, and not invertebrate pests, such as mosquitoes and termites. Endemic invertebrates are likely to represent a portion of the coqui diet, but it remains uncertain which endemic invertebrates are most threatened by coqui predation and whether there will be indirect effects that benefit or harm them. It could be predicted that coquis would compete with endemic birds for invertebrate prey, but there is presently little overlap in the habitats used by coquis and endemic birds. Although, coquis may make bird re‐invasion into lowland ecosystems more difficult; alternatively, coquis could serve as an additional food source for some endemic birds. Finally, it could be predicted that coquis serve as a food source for endemic‐bird predators, such as rats and mongoose, and bolster their abundance. Preliminary data suggest that coquis will not bolster rat or mongoose populations. Managing coqui populations in Hawaii has been a challenge. A population has not yet been eradicated using citric acid, the only federally approved pesticide for coquis. It is unlikely that coquis will ever be eradicated from the islands of Hawaii and Maui, where there are now hundreds of populations. Quick and severe responses to new introductions may be the only effective means of containing the spread of the coqui.     

Different prey resources suggest little competition between non-native frogs and insectivorous birds despite isotopic niche overlap

Non-native amphibians often compete with native amphibians in their introduced range, but their competitive effects on other vertebrates are less well known. The Puerto Rican coqui frog (Eleutherodactylus coqui) has colonized the island of Hawaii, and has been hypothesized to compete with insectivorous birds and bats. To address if the coqui could compete with these vertebrates, we used stable isotope analyses to compare the trophic position and isotopic niche overlap between the coqui, three insectivorous bird species, and the Hawaiian hoary bat. Coquis shared similar trophic position to Hawaii amakihi, Japanese white-eye, and red-billed leiothrix. Coquis were about 3 ‰ less enriched in δ¹⁵N than the Hawaiian hoary bat, suggesting the bats feed at a higher trophic level than coquis. Analyses of potential diet sources between coquis and each of the three bird species indicate that there was more dietary overlap between bird species than any of the birds and the coqui. Results suggest that Acari, Amphipoda, and Blattodea made up >90% of coqui diet, while Araneae made up only 2% of coqui diet, but approximately 25% of amakihi and white-eye diet. The three bird species shared similar proportions of Lepidoptera larvae, which were ~25% of their diet. Results suggest that coquis share few food resources with insectivorous birds, but occupy a similar trophic position, which could indicate weak competition. However, resource competition may not be the only way coquis impact insectivorous birds, and future research should examine whether coqui invasions are associated with changes in bird abundance.     

Parasite loss and introduced species: a comparison of the parasites of the Puerto Rican tree frog, (<Emphasis Type="Italic">Eleutherodactylus coqui</Emphasis>), in its native and introduced ranges

The Puerto Rican frog, Eleutherodactylus coqui has invaded Hawaii and reached densities far exceeding those in their native range. One possible explanation for the success of E. coqui in its introduced range is that it lost its co-evolved parasites in the process of the invasion. We compared the parasites of E. coqui in its native versus introduced range. We collected parasite data on 160 individual coqui frogs collected during January-April 2006 from eight populations in Puerto Rico and Hawaii. Puerto Rican coqui frogs had higher species richness of parasites than Hawaiian coqui frogs. Parasite prevalence and intensity were significantly higher in Hawaii, however this was likely a product of the life history of the dominant parasite and its minimal harm to the host. This suggests that the scarcity of parasites may be a factor contributing to the success of Eleutherodactylus coqui in Hawaii.     

Predicting the distribution potential of an invasive frog using remotely sensed data in Hawaii

Aim Eleutherodactylus coqui (commonly known as the coqui) is a frog species native to Puerto Rico and non‐native in Hawaii. Despite its ecological and economic impacts, its potential range in Hawaii is unknown, making control and management efforts difficult. Here, we predicted the distribution potential of the coqui on the island of Hawaii. Location Puerto Rico and Hawaii. Methods We predicted its potential distribution in Hawaii using five biophysical variables derived from Moderate Resolution Imaging Spectroradiometer (MODIS) as predictors, presence/absence data collected from Puerto Rico and Hawaii and three classification methods – Classification Trees (CT), Random Forests (RF) and Support Vector Machines (SVM). Results Models developed separately using data from the native range and the invaded range predicted potential coqui habitats in Hawaii with high performance. Across the three classification methods, mean area under the ROC curve (AUC) was 0.75 for models trained using the native range data and 0.88 for models trained using the invaded range data. We achieved the highest AUC value of 0.90 using RF for models trained with invaded range data. Main conclusions Our results showed that the potential distribution of coquis on the island of Hawaii is much larger than its current distribution, with RF predicting up to 49% of the island as suitable coqui habitat. Predictions also show that most areas with an elevation between 0 and 2000 m are suitable coqui habitats, whereas the cool and dry high elevation areas beyond 2000 m elevation are unsuitable. Results show that MODIS‐derived biophysical variables are capable of characterizing coqui habitats in Hawaii.     

The effect of aquatic and terrestrial habitat characteristics on occurrence and breeding probability in a montane amphibian: insights from a spatially explicit multistate occupancy model

Understanding species distribution and predicting range shifts are major goals of ecology and biogeography. Obtaining reliable predictions of how species distribution might change in response to habitat change requires knowledge of habitat availability, occupancy, use for breeding, and spatial autocorrelation in these parameters. Amphibians in alpine areas provide an excellent model system for disentangling habitat drivers of occupancy from that of breeding while explicitly accounting for spatial autocorrelation. We focused on the widespread common frog (Rana temporaria) inhabiting alpine lakes in the Southern Carpathians, Romania. We used single season multistate occupancy models developed to account for imperfect detection and spatial autocorrelation to estimate the occupancy and breeding probabilities and to evaluate their response to habitat characteristics. We found that frogs do not occur in all water bodies [occupancy probability: 0.697; 95% credible interval (0.614, 0.729)] and do not breed in a substantial proportion of water bodies where they occur [breeding probability conditional on occupancy: 0.707; 95% credible interval (0.670, 0.729)]. Habitat characteristics explain water body occupancy but not breeding probability; and altitude, water body surface area, water body sinuosity and permanency, presence of invertebrates, and grazing along the banks all had positive effects on occupancy. We also detected strong spatial autocorrelation in occupancy and breeding probabilities. Thus, our results indicate that habitat choice by montane amphibians is influenced by both spatial autocorrelation and habitat characteristics. Because spatial autocorrelations matter and because the presence of adults is not the same as the presence of a reproducing population, it will be difficult to predict the effects of habitat change on high altitude amphibian populations.     

Calls of Recently Introduced Coquí Frogs Do Not Interfere with Cricket Phonotaxis in Hawaii

Acoustically-signaling animals such as crickets may experience interference from environmental noise, a particular concern given the rise in anthropogenic or other novel sources of sound. We examined the potential for acoustic interference of female phonotaxis to calling song in the Pacific field cricket (Teleogryllus oceanicus) by invasive coqui frogs (Eleutherodactylus coqui) in Hawaii. The frogs were introduced to Hawaii from Puerto Rico in the 1980s. When female crickets were exposed to male calling songs with and without simultaneous broadcast of a coqui chorus, they were equally likely to move toward the cricket song, regardless of the location of the frog sound (ground level or above ground). Unlike some species of frogs and birds, T. oceanicus do not appear to experience acoustic interference from an introduced signaler, even though the introduced species’ calls subjectively seem to be masking the crickets’ songs.     

Estimating density of calling male Eleutherodactylus coqui in Hawaii from audio recordings of the nighttime frog chorus

Analysis of sound pressure data obtained from audio recordings of night-time Eleutherodactylus coqui chorusing was used to estimate population density of calling male coqui frogs in the major Hawaiian island populations. Study plots 400 m² in size (20 by 20 m; n = 32) were established on the islands of Hawaii and Maui to provide venues for field counting calling male frogs to investigate the ground truth relationship with the recorded sound pressure data. Male coqui frogs emit a two-note ‘co-qui’ advertisement call at regular intervals. Exploiting the temperature-related and relatively consistent male coqui call repetition rate and the average emitted ‘co’ call sound pressure level (70.4 dB at 1 m), cue counts obtained from a correlation process available in the program Raven were also used to estimate the density of calling male coqui. The study methodology included comparing linear regression results between the field-counted frogs and the number of frogs obtained from the cue count method. Both methods revealed a linear relationship between calling male population density and associated average sound pressure of the resulting chorus.     

Growth and uptake of mineral by embryos of the direct-developing frog Eleutherodactylus coqui

Embryos of the direct-developing frog Eleutherodactylus coqui take up small quantities of yolk and yolk mineral early in incubation but increase their uptake of yolk reserves at later stages of development. Growth and accumulation of calcium and magnesium by embryos also occur slowly at first and at a higher rate later. Accumulation of calcium and magnesium by embryos is largely a function of variation in size of embryos, but uptake of phosphorus is unrelated to size. Althrough patterns of growth and uptake of mineral by embryonic coquis resemble those for embryos of oviparous amniotes, embryonic coquis do not deplete the yolk of its nutrients to the same degree. Thus, residual yolk of coqui hatchlings contains a high percentage of the nutrient reserves originally present in the egg. This difference between embryonic coquis and embryos of oviparous amniotes may indicate that transfer of nutrients from yolk to embryo becomes limiting during the grwoth phase. Alternatively, some aspects of the neurologic system are so poorly developed at hatching that coqui may not be able to find prey effectively. A large nutrient reserve could sustain hatchling while the neurologic system continues to mature.     

Estimating site occupancy rates for aquatic plants using spatial sub-sampling designs when detection probabilities are less than one

Estimation of site occupancy rates when detection probabilities are <1 is well established in wildlife science. Data from multiple visits to a sample of sites are used to estimate detection probabilities and the proportion of sites occupied by focal species. In this article we describe how site occupancy methods can be applied to estimate occupancy rates of plants and other sessile organisms. We illustrate this approach and the pitfalls of ignoring incomplete detection using spatial data for 2 aquatic vascular plants collected under the Upper Mississippi River's Long Term Resource Monitoring Program (LTRMP). Site occupancy models considered include: a naïve model that ignores incomplete detection, a simple site occupancy model assuming a constant occupancy rate and a constant probability of detection across sites, several models that allow site occupancy rates and probabilities of detection to vary with habitat characteristics, and mixture models that allow for unexplained variation in detection probabilities. We used information theoretic methods to rank competing models and bootstrapping to evaluate the goodness-of-fit of the final models. Results of our analysis confirm that ignoring incomplete detection can result in biased estimates of occupancy rates. Estimates of site occupancy rates for 2 aquatic plant species were 19–36% higher compared to naive estimates that ignored probabilities of detection <1. Simulations indicate that final models have little bias when 50 or more sites are sampled, and little gains in precision could be expected for sample sizes >300. We recommend applying site occupancy methods for monitoring presence of aquatic species.     

Calling at the highway: The spatiotemporal constraint of road noise on Pacific chorus frog communication

Loss of acoustic habitat due to anthropogenic noise is a key environmental stressor for vocal amphibian species, a taxonomic group that is experiencing global population declines. The Pacific chorus frog (Pseudacris regilla) is the most common vocal species of the Pacific Northwest and can occupy human‐dominated habitat types, including agricultural and urban wetlands. This species is exposed to anthropogenic noise, which can interfere with vocalizations during the breeding season. We hypothesized that Pacific chorus frogs would alter the spatial and temporal structure of their breeding vocalizations in response to road noise, a widespread anthropogenic stressor. We compared Pacific chorus frog call structure and ambient road noise levels along a gradient of road noise exposures in the Willamette Valley, Oregon, USA. We used both passive acoustic monitoring and directional recordings to determine source level (i.e., amplitude or volume), dominant frequency (i.e., pitch), call duration, and call rate of individual frogs and to quantify ambient road noise levels. Pacific chorus frogs were unable to change their vocalizations to compensate for road noise. A model of the active space and time (“spatiotemporal communication”) over which a Pacific chorus frog vocalization could be heard revealed that in high‐noise habitats, spatiotemporal communication was drastically reduced for an individual. This may have implications for the reproductive success of this species, which relies on specific call repertoires to portray relative fitness and attract mates. Using the acoustic call parameters defined by this study (frequency, source level, call rate, and call duration), we developed a simplified model of acoustic communication space–time for this species. This model can be used in combination with models that determine the insertion loss for various acoustic barriers to define the impact of anthropogenic noise on the radius of communication in threatened species. Additionally, this model can be applied to other vocal taxonomic groups provided the necessary acoustic parameters are determined, including the frequency parameters and perception thresholds. Reduction in acoustic habitat by anthropogenic noise may emerge as a compounding environmental stressor for an already sensitive taxonomic group.     

Human-mediated Escalation of a Formerly Eradicable Problem: The Invasion of Caribbean Frogs in the Hawaiian Islands

Two species of neotropical frog, Eleutherodactylus coqui and E. planirostris, have been introduced into the State of Hawaii via the horticulture trade. E. coqui was introduced prior to 1988 and E. planirostris was first reported in 1994. Since these dates frog colonies have rapidly spread accidentally and intentionally and frog abundance within colonies has grown rapidly. Although these frogs were originally restricted to horticulture sites, they are now found in residential areas, resorts and hotels, and public lands. Due to the high potential biomass of introduced frogs there are realistic anthropogenic and ecological concerns associated with the spread of these frogs. Though there currently is a tool that can be used for localized control of frogs in limited circumstances, overall efforts by Federal, State, and County agencies to control the frog in Hawaii have been hampered by limited authorities and funds, disbelief in the threat, and the reluctance to act.     

When Is a Species Declining? Optimizing Survey Effort to Detect Population Changes in Reptiles

Biodiversity monitoring programs need to be designed so that population changes can be detected reliably. This can be problematical for species that are cryptic and have imperfect detection. We used occupancy modeling and power analysis to optimize the survey design for reptile monitoring programs in the UK. Surveys were carried out six times a year in 2009-2010 at multiple sites. Four out of the six species - grass snake, adder, common lizard, slow-worm -were encountered during every survey from March-September. The exceptions were the two rarest species -- sand lizard and smooth snake - which were not encountered in July 2009 and March 2010 respectively. The most frequently encountered and most easily detected species was the slow-worm. For the four widespread reptile species in the UK, three to four survey visits that used a combination of directed transect walks and artificial cover objects resulted in 95% certainty that a species would be detected if present. Using artificial cover objects was an effective detection method for most species, considerably increased the detection rate of some, and reduced misidentifications. To achieve an 85% power to detect a decline in any of the four widespread species when the true decline is 15%, three surveys at a total of 886 sampling sites, or four surveys at a total of 688 sites would be required. The sampling effort needed reduces to 212 sites surveyed three times, or 167 sites surveyed four times, if the target is to detect a true decline of 30% with the same power. The results obtained can be used to refine reptile survey protocols in the UK and elsewhere. On a wider scale, the occupancy study design approach can be used to optimize survey effort and help set targets for conservation outcomes for regional or national biodiversity assessments.     

小气候对东北林蛙生长发育和繁殖的影响

于2009年5月—2010年5月,通过分组实验,研究了在大棚增温环境和围栏常温模拟林下环境条件下东北林蛙(Rana dybowskii)的生长发育情况。结果表明:与围栏常温模拟林下条件相比,大棚圈舍内平均气温、地温和湿度显著升高,有效积温增加;林蛙发育历期缩短,性成熟提前,二龄雌蛙性成熟比例增加,大棚圈舍的雌蛙的繁殖率为93.1%,围栏圈舍雌蛙的繁殖率为76.0%;在大棚中林蛙生长速度加快,饲养的一、二龄林蛙体重分别达到(6.30±2.62)g和(36.55±11.79)g,较围栏中体重提高了85.5%和56.6%;在大棚里林蛙出蛰时间、进食时间提前,停食时间、入蛰时间延后,生长期间延长,每窝平均产卵量增加,林蛙体重与产卵量呈显著的正相关(r=0.921,P<0.01)。     

High level of cryptic species diversity revealed by sympatric lineages of Southeast Asian forest frogs

Amphibians tend to exhibit conservative morphological evolution, and the application of molecular and bioacoustic tools in systematic studies have been effective at revealing morphologically 'cryptic' species within taxa that were previously considered to be a single species. We report molecular genetic findings on two forest-dwelling ranid frogs from localities across Southeast Asia, and show that sympatric evolutionary lineages of morphologically cryptic frogs are a common pattern. These findings imply that species diversity of Southeast Asian frogs remains significantly underestimated, and taken in concert with other molecular investigations, suggest there may not be any geographically widespread, forest-dwelling frog species in the region. Accurate assessments of diversity and distributions are needed to mitigate extinctions of evolutionary lineages in these threatened vertebrates.     

Ecology and taxonomy-driven deviations in the frog call–body size relationship across the diverse Australian frog fauna

Relationships between some properties of frog calls and body size are widely recognized. However, generality across call components and diverse faunas, and sources of deviation, remain poorly tested. Using 116 east Australian frog species, we tested the relationship between three call traits and body size, and the effects of taxonomic family and calling habitat. Call dominant frequency (DF) has a highly significant negative relationship with size, whereas call duration and pulse rate do not. Frog families show the same slope of relationship between DF and size, but hylids call at significantly higher frequency relative to size. Within hylids, stream breeders call at significantly lower DF than pool breeders of comparable size – below the DF of stream noise in typical breeding habitat – a shift likely to enhance signal detection against background environmental noise. This contrasts with all previous observations from other regions that frogs call at high (even ultrasonic) frequency to avoid masking by stream noise.     

Assessing breeding success in common woodland birds using a novel method

Capsule Repeated counts of fledged broods can provide a useful estimate of breeding success for most common woodland birds.

Aims To assess the efficacy of comparing fledged-brood survey data with territory mapping using simple mark–recapture analysis techniques to provide an estimate of breeding success for common woodland birds that does not involve finding nests.

Methods Three observers undertook territory mapping surveys of adults, followed by counts of fledged broods four times a week during May–July 2007 in two 15 ha woods each, both in southern England. Using known fledging to maturity periods, these counts were used to calculate daily detection probabilities for broods of ubiquitous species. These enabled fledged brood territory occupancy probabilities (i.e. brood to territory ratios) to be estimated that take account of the possibility that broods were present but missed by surveys.

Results Of the 19 species found in all six woods, mean daily detection probability estimates for fledged broods of 17 species ranged from 0.17 to 0.50 with significant variation between woods for 12 species, but within region/observer for four species. The mean probability of detecting a brood at least once was over 75% using four visits per week and over 50% using two visits. Only for Great Spotted Woodpeckers Dendrocopos major and Garden Warblers Sylvia borin was the fledging period too short and the daily detection probability too low to provide a reasonable estimate of the territory occupancy probability.

Conclusion Daily detection probabilities for fledged broods of most common woodland birds were sufficiently high to enable useable estimates of fledged-brood territory occupancy probabilities to be made based on a survey programme involving two or three visits per week between late May and the end June. The method used may have application as a means of providing a relatively easily derived productivity index for woodland bird monitoring programmes or for research studies.     

Rare but not lost: Endemic mountain lizard occupancy following megafire and grazing disturbances

Wildfire and grazing by invasive herbivores can influence habitat suitability for ground-dwelling fauna, such as reptiles. Australia has a large and diverse reptile fauna, with the Australian Alps bioregion in the southeast of the continent supporting a disproportionately high number of threatened species. In this bioregion, many species are threatened by fire, habitat loss or modification, and invasive species. The range of one such threatened endemic lizard, Cyclodomorphus praealtus (family Scincidae), was impacted by the 2019–20 megafires and is also subject to widespread grazing by invasive species. We investigated the relationship between C. praealtus site occupancy and fire and grazing. We completed 2045 surveys across 120 sites over 4 years, detecting the species at 43% of sites and increasing the species' known geographic range. Using single season detection occupancy models, we found C. praealtus occupancy was not associated with elevation, vegetation height or whether the site was burnt, but was positively associated with grazing activity. Our results indicate that C. praealtus has the capacity to persist following a single fire in some cases, and that habitats with high occupancy probabilities are subject to high grazing pressure. However, our results do not rule out more nuanced impacts associated with these disturbances, which affect a large proportion of C. praealtus' habitat. Our cumulative detection probability calculations revealed that considerable survey effort is often required to determine C. praealtus site occupancy. We therefore recommend that impact assessments assume species presence within areas of suitable habitat within the species' range. Our study improves our understanding of disturbance impacts on C. praealtus' occupancy, while demonstrating the need for sufficiently resourced impact assessments for cryptic and threatened species.     

Red-Shouldered Hawk Occupancy Surveys in Central Minnesota,USA

ABSTRACT Forest-dwelling raptors are often difficult to detect because many species occur at low density or are secretive. Broadcasting conspecific vocalizations can increase the probability of detecting forest-dwelling raptors and has been shown to be an effective method for locating raptors and assessing their relative abundance. Recent advances in statistical techniques based on presence—absence data use probabilistic arguments to derive probability of detection when it is < 1 and to provide a model and likelihood-based method for estimating proportion of sites occupied. We used these maximum-likelihood models with data from red-shouldered hawk (Buteo lineatus) call-broadcast surveys conducted in central Minnesota, USA, in 1994–1995 and 2004–2005. Our objectives were to obtain estimates of occupancy and detection probability 1) over multiple sampling seasons (yr), 2) incorporating within-season time-specific detection probabilities, 3) with call type and breeding stage included as covariates in models of probability of detection, and 4) with different sampling strategies. We visited individual survey locations 2–9 times per year, and estimates of both probability of detection (range = 0.28-0.54) and site occupancy (range = 0.81-0.97) varied among years. Detection probability was affected by inclusion of a within-season time-specific covariate, call type, and breeding stage. In 2004 and 2005 we used survey results to assess the effect that number of sample locations, double sampling, and discontinued sampling had on parameter estimates. We found that estimates of probability of detection and proportion of sites occupied were similar across different sampling strategies, and we suggest ways to reduce sampling effort in a monitoring program.     

Parallel microtubules and other conserved elements of dorsal axial specification in the direct developing frog, <Emphasis Type="Italic">Eleutherodactylus coqui</Emphasis>

Specification of the dorsal axis in commonly studied frogs, such as Xenopus laevis and Rana pipiens, depends on a microtubule-mediated movement of cytoplasm in the fertilized egg. The Puerto Rican tree frog, Eleutherodactylus coqui, has an egg that is twenty times the volume of that of X. laevis, raising the question as to whether the mechanism of dorsal axial specification is conserved in these large eggs. Fertilized eggs of E. coqui develop a transient array of parallel microtubules, similar to other frogs, but proportionately larger. The array persists after first cleavage, longer than in other frogs, and is gone by the third cleavage. Correlated with the longer life of the parallel microtubules, both 2- and 8-cell E. coqui embryos remain sensitive to gravity-mediated axial specification, a sensitivity lost in X. laevis before the 2-cell stage. Activation of the Wnt/beta-catenin pathway by injected Xwnt8 RNA causes axial formation as in X. laevis. The results indicate that elements of dorsal axial specification are conserved in E. coqui, but they occur later compared to in X. laevis.