Is habitat enhancement a viable strategy for conserving New Zealand's endemic lizards?

<p>In our current era, the Anthropocene, species are disappearing at an unprecedented rate due to the impact of humans on Earth’s environments. Of the many causes of these extinctions, habitat loss is thought to be the most severe. Three habitat management strategies are available for halting habitat loss: reservation, restoration and reconciliation. The latter two of these strategies actively seek to improve the ability of degraded or lost habitats to support species. If successful on a large enough scale, use of restoration and reconciliation (hereafter referred to collectively as ‘habitat enhancement’) could reverse the effects of habitat loss. I evaluated the viability of habitat enhancement for the conservation of New Zealand’s lizard fauna. 83% of New Zealand’s 106+ endemic species are threatened or at risk of extinction. While habitat loss is one key driver of declines, predation by invasive mammals is the other. Neither of these processes are well understood. Habitat enhancement is increasingly being employed in New Zealand by landowners, community groups, conservationists, and businesses as a strategy for mitigating lizard declines, but outcomes are rarely investigated comprehensively. This is concerning because habitat manipulation potentially affects both exotic and native species, which has led to unexpected negative effects on threatened fauna in New Zealand and overseas. I posed four questions to help address this knowledge gap. (1) What habitat enhancement strategies are available for reptiles, and have they produced successful conservation outcomes? (2) How do habitat characteristics affect populations and communities of endemic New Zealand lizards? (3) How does the presence of invasive mammals affect populations and communities of endemic New Zealand lizards over intermediate to long-term time frames? (4) Can habitat enhancement produce positive conservation outcomes in the presence of invasive mammals? A review of the global literature on habitat enhancement for reptiles identified 75 studies documenting 577 responses of 251 reptile species. For outcome evaluation, I adapted an existing stage-based framework for assessment of translocation success. High levels of success (84-85%) at Stages 1 (use of enhanced habitat) and 2 (evidence of reproduction in enhanced habitat) suggested that enhancement could be useful for creating areas that can be inhabited, and reproduced in, by reptiles. Fewer cases were successful at Stage 3 (30%; improvement of at least one demographic parameter demonstrated in enhanced habitat) or Stage 4 (43%; self-sustaining or source population established in enhanced habitat). Additionally, only 1% of the 577 cases sufficiently examined or modelled long-term population trends to allow evaluation against the Stage 4 criterion. Thus, there was a lack of evidence indicating that enhancement could result in higher population growth rates, or reduced extinction risk, of reptiles. I conducted field work in the Wellington region to investigate the effects of habitat characteristics and mammals on terrestrial lizards inhabiting coastal environments. Surveys conducted in two mammal-invaded mainland areas and on two mammal-free offshore islands showed that presence or absence of invasive mammals had a stronger effect on lizard community structure than habitat variables. However, occupancy probabilities of northern grass skinks Oligosoma polychroma and Raukawa geckos Woodworthia maculata were positively correlated with increasing cover of divaricating shrubs. O. polychroma were also more likely to occupy patches with increasing cover by non-Muehlenbeckia vines. Mark-recapture studies were conducted at two mammal-invaded mainland sites to investigate the current abundance of lizard species: Turakirae Head and Pukerua Bay. Estimated densities of O. polychroma ranged between 3,980 and 4,078 individuals / ha and W. maculata between 4,067 and 38,372 individuals / ha. Other species known to occur, at least historically, at each site were either not detected or comprised only a small proportion of total lizard captures. Analysis of longitudinal lizard monitoring data available for Pukerua Bay, Turakirae Head, and an additional mammal-invaded site, Baring Head, did not reveal a significant decline in abundance, occupancy, or catch rates of O. polychroma over time periods ranging between six and 34 years, nor of W. maculata over six to 49 years. Habitat information available for Baring Head showed that the probability of local extinction of W. maculata was significantly lower at rocky sites. Finally, I conducted a before-after-control-impact habitat enhancement experiment on lizard communities inhabiting 100 m2 plots on the mammal-invaded Miramar Peninsula. After a six-month pre-enhancement monitoring period, native plants and gravel piles were added to enhancement plots and lizard monitoring continued for a further nine months. Enhancement did not significantly affect plot use, body condition, or evidence of reproduction in Oligosoma aeneum, O. polychroma or W. maculata, but were considered successful at Stages 1 and 2 due to the absence of a negative effect. Neither the abundance, probability of entry into plots by birth or immigration, nor apparent survival of O. aeneum was significantly affected by enhancement (Stage 3). Apparent survival of O. polychroma increased significantly in response to enhancement, but this did not result in increased abundance. Adding gravel and native vegetation (especially divaricating shrubs and vines) may be a suitable strategy for creating habitat in invaded coastal landscapes for O. polychroma and W. maculata. However, most of the other lizard species that would have historically occurred in mammal-invaded coastal areas of Wellington appeared to be sensitive to sustained mammal presence, even with low-to-moderate levels of control in operation. Therefore, habitat enhancement without intensive mammal control or eradication is not expected to benefit these species, nor be capable of restoring coastal lizard communities. In invaded landscapes it is, at best, a reconciliation measure that could allow co-existence of an endemic lizard community comprised of common species with invasive mammals. However, habitat enhancement could still be useful for restoring lizard communities in mammal-free sanctuaries.</p>

Is habitat enhancement a viable strategy for conserving New Zealand's endemic lizards?

<p>In our current era, the Anthropocene, species are disappearing at an unprecedented rate due to the impact of humans on Earth’s environments. Of the many causes of these extinctions, habitat loss is thought to be the most severe. Three habitat management strategies are available for halting habitat loss: reservation, restoration and reconciliation. The latter two of these strategies actively seek to improve the ability of degraded or lost habitats to support species. If successful on a large enough scale, use of restoration and reconciliation (hereafter referred to collectively as ‘habitat enhancement’) could reverse the effects of habitat loss. I evaluated the viability of habitat enhancement for the conservation of New Zealand’s lizard fauna. 83% of New Zealand’s 106+ endemic species are threatened or at risk of extinction. While habitat loss is one key driver of declines, predation by invasive mammals is the other. Neither of these processes are well understood. Habitat enhancement is increasingly being employed in New Zealand by landowners, community groups, conservationists, and businesses as a strategy for mitigating lizard declines, but outcomes are rarely investigated comprehensively. This is concerning because habitat manipulation potentially affects both exotic and native species, which has led to unexpected negative effects on threatened fauna in New Zealand and overseas. I posed four questions to help address this knowledge gap. (1) What habitat enhancement strategies are available for reptiles, and have they produced successful conservation outcomes? (2) How do habitat characteristics affect populations and communities of endemic New Zealand lizards? (3) How does the presence of invasive mammals affect populations and communities of endemic New Zealand lizards over intermediate to long-term time frames? (4) Can habitat enhancement produce positive conservation outcomes in the presence of invasive mammals? A review of the global literature on habitat enhancement for reptiles identified 75 studies documenting 577 responses of 251 reptile species. For outcome evaluation, I adapted an existing stage-based framework for assessment of translocation success. High levels of success (84-85%) at Stages 1 (use of enhanced habitat) and 2 (evidence of reproduction in enhanced habitat) suggested that enhancement could be useful for creating areas that can be inhabited, and reproduced in, by reptiles. Fewer cases were successful at Stage 3 (30%; improvement of at least one demographic parameter demonstrated in enhanced habitat) or Stage 4 (43%; self-sustaining or source population established in enhanced habitat). Additionally, only 1% of the 577 cases sufficiently examined or modelled long-term population trends to allow evaluation against the Stage 4 criterion. Thus, there was a lack of evidence indicating that enhancement could result in higher population growth rates, or reduced extinction risk, of reptiles. I conducted field work in the Wellington region to investigate the effects of habitat characteristics and mammals on terrestrial lizards inhabiting coastal environments. Surveys conducted in two mammal-invaded mainland areas and on two mammal-free offshore islands showed that presence or absence of invasive mammals had a stronger effect on lizard community structure than habitat variables. However, occupancy probabilities of northern grass skinks Oligosoma polychroma and Raukawa geckos Woodworthia maculata were positively correlated with increasing cover of divaricating shrubs. O. polychroma were also more likely to occupy patches with increasing cover by non-Muehlenbeckia vines. Mark-recapture studies were conducted at two mammal-invaded mainland sites to investigate the current abundance of lizard species: Turakirae Head and Pukerua Bay. Estimated densities of O. polychroma ranged between 3,980 and 4,078 individuals / ha and W. maculata between 4,067 and 38,372 individuals / ha. Other species known to occur, at least historically, at each site were either not detected or comprised only a small proportion of total lizard captures. Analysis of longitudinal lizard monitoring data available for Pukerua Bay, Turakirae Head, and an additional mammal-invaded site, Baring Head, did not reveal a significant decline in abundance, occupancy, or catch rates of O. polychroma over time periods ranging between six and 34 years, nor of W. maculata over six to 49 years. Habitat information available for Baring Head showed that the probability of local extinction of W. maculata was significantly lower at rocky sites. Finally, I conducted a before-after-control-impact habitat enhancement experiment on lizard communities inhabiting 100 m2 plots on the mammal-invaded Miramar Peninsula. After a six-month pre-enhancement monitoring period, native plants and gravel piles were added to enhancement plots and lizard monitoring continued for a further nine months. Enhancement did not significantly affect plot use, body condition, or evidence of reproduction in Oligosoma aeneum, O. polychroma or W. maculata, but were considered successful at Stages 1 and 2 due to the absence of a negative effect. Neither the abundance, probability of entry into plots by birth or immigration, nor apparent survival of O. aeneum was significantly affected by enhancement (Stage 3). Apparent survival of O. polychroma increased significantly in response to enhancement, but this did not result in increased abundance. Adding gravel and native vegetation (especially divaricating shrubs and vines) may be a suitable strategy for creating habitat in invaded coastal landscapes for O. polychroma and W. maculata. However, most of the other lizard species that would have historically occurred in mammal-invaded coastal areas of Wellington appeared to be sensitive to sustained mammal presence, even with low-to-moderate levels of control in operation. Therefore, habitat enhancement without intensive mammal control or eradication is not expected to benefit these species, nor be capable of restoring coastal lizard communities. In invaded landscapes it is, at best, a reconciliation measure that could allow co-existence of an endemic lizard community comprised of common species with invasive mammals. However, habitat enhancement could still be useful for restoring lizard communities in mammal-free sanctuaries.</p>

Age-specific survival in an English Twite population.

Like many bird species associated with agricultural habitats in the UK, the Twite Linaria flavirostris has undergone severe declines over recent decades due to habitat degradation, with populations in England, Wales and Ireland now restricted to a few small pockets. However, the demographic drivers of these declines are still largely unresolved. We estimated the survival of Twite from a small population at the southernmost edge of the English range in Derbyshire using capture-mark-recapture data from 2016–2019. Annual apparent survival for juveniles (0.14–0.34) was lower than for adults (0.29–0.56) and less than that of other Cardueline finches. Our results suggest that low juvenile survival may be one demographic driver underpinning the recent decline of the Derbyshire Twite population, although we also cannot rule out the possibility that differences in emigration of juveniles and adults from the population also contribute to the observed age-specific apparent survival rates.

Strong evidence supporting a relationship between colour pattern and apparent survival in common crossbills

Carotenoid staining has been repeatedly shown to serve as a sexually selected individual quality signal. In different species, individuals that show brighter carotenoid-based signals have been found to have superior feeding abilities, recover faster from disease, and generally enjoy better body condition. In the common crossbill (Loxia curvirostra), the colour has also been related to the different populations, with northern and central European populations being described as redder than those in the Mediterranean region. A study in the Pyrenees showed that long-winged individuals had lower apparent survival, and the proportion of red individuals was higher in long-winged birds, concluding that they could be nomadic birds (that travel long distances). A priori, if the red crossbills are more mobile than the yellow and orange ones, their apparent survival will be lower. However, in our study, red males showed a greater survival than males of other colours and almost double than that of the yellow ones. These results suggest that red coloration is linked to higher quality individuals regardless of their mobility.

High turn-over rates at the upper range limit and elevational source-sink dynamics in a widespread songbird

The formation of an upper distributional range limit for species breeding along mountain slopes is often based on environmental gradients resulting in changing demographic rates towards high elevations. However, we still lack an empirical understanding of how the interplay of demographic parameters forms the upper range limit in highly mobile species. Here, we study apparent survival and within-study area dispersal over a 700 m elevational gradient in barn swallows (Hirundo rustica) by using 15 years of capture-mark-recapture data. Annual apparent survival of adult breeding birds decreased while breeding dispersal probability of adult females, but not males increased towards the upper range limit. Individuals at high elevations dispersed to farms situated at elevations lower than would be expected by random dispersal. These results suggest higher turn-over rates of breeding individuals at high elevations, an elevational increase in immigration and thus, within-population source-sink dynamics between low and high elevations. The formation of the upper range limit therefore is based on preference for low-elevation breeding sites and immigration to high elevations. Thus, shifts of the upper range limit are not only affected by changes in the quality of high-elevation habitats but also by factors affecting the number of immigrants produced at low elevations.

What do we know about survival of Common cranes? An elementary introduction with Euring databank

The increase of the western populations of Common cranes (Grus grus) in the last five decades highlights the need to estimate survival rates. According to Euring databank (EDB), the oldest Common crane ever known was 27 years old in year 2017. This lifespan was obtained by means of 24,900 recoveries of 2,124 ringed cranes collected between years 1936 and 2017. Nearly all cranes were ringed and observed in the last 30 years, and therefore the elapsed time was not enough to reach the maximum longevity reported for the species in captivity (43 years, Mitchell 1911). Life expectancy was five years on average after the ring was attached. Here we provide some elementary analyses to calculate the annual apparent survival rate (ϕ = 0.85) and the annual encounter probability (p = 0.45) of Common cranes, as a first step to advance in the knowledge of the species' population dynamics. The great increase of breeding and wintering crane populations in western Europe in the last decades remains largely unexplained.

Leopard Panthera pardus density and survival in an ecosystem with depressed abundance of prey and dominant competitors

The leopard Panthera pardus is in range-wide decline, and many populations are highly threatened. Prey depletion is a major cause of global carnivore declines, but the response of leopard survival and density to this threat is unclear: by reducing the density of a dominant competitor (the lion Panthera leo) prey depletion could create both costs and benefits for subordinate competitors. We used capture–recapture models fitted to data from a 7-year camera-trap study in Kafue National Park, Zambia, to obtain baseline estimates of leopard population density and sex-specific apparent survival rates. Kafue is affected by prey depletion, and densities of large herbivores preferred by lions have declined more than the densities of smaller herbivores preferred by leopards. Lion density is consequently low. Estimates of leopard density were comparable to ecosystems with more intensive protection and favourable prey densities. However, our study site is located in an area with good ecological conditions and high levels of protection relative to other portions of the ecosystem, so extrapolating our estimates across the Park or into adjacent Game Management Areas would not be valid. Our results show that leopard density and survival within north-central Kafue remain good despite prey depletion, perhaps because (1) prey depletion has had weaker effects on preferred leopard prey compared to larger prey preferred by lions, and (2) the density of dominant competitors is consequently low. Our results show that the effects of prey depletion can be more complex than uniform decline of all large carnivore species, and warrant further investigation.

Emigration Effects on Estimates of Age- and Sex-specific Survival of Small Mammals

Age- and sex-specific survival estimates are crucial to understanding important life-history characteristics and variation in these estimates can be a key driver of population dynamics. When estimating survival using Cormack-Jolly-Seber (CJS) models and capture-recapture data, emigration is typically assumed to have a negligible effect on estimates such that apparent survival is indistinguishable from true survival. Consequently, especially for populations or age classes with high dispersal rates, apparent survival estimates are often biased low and temporal patterns in survival might be masked when site fidelity varies temporally. We used 9 years of annual mark-recapture data to estimate age-, sex-, and time-specific apparent survival of Humboldt's flying squirrels (Glaucomys oregonensis) and Townsend's chipmunks (Neotamias townsendii). For Humboldt's flying squirrels, these estimates support a small body of research investigating potential variation of survival among age and sex classes, but age- and sex-specific survival has not been evaluated for Townsend's chipmunks. We also quantified the effects of age- and sex-specific emigration on confounded estimates of apparent survival. Our estimates of juvenile flying squirrel survival were high relative to other small mammal species and estimates for both species were variable among years. We found survival differed moderately among age and sex classes for Humboldt's flying squirrels, but little among age and sex classes for Townsend's chipmunks, and that the degree to which emigration confounded apparent survival estimates varied substantially among years. Without correcting for emigration, apparent survival estimates were lower and temporal variation was obscured, particularly for male Humboldt's flying squirrels and female Townsend's chipmunks. Our results demonstrate that emigration can influence commonly used estimates of apparent survival. Unadjusted estimates confounded the interpretation of differences in survival between age and sex classes and masked potential temporal patterns in survival because the magnitude of adjustment varied among years. We conclude that apparent survival estimators are robust during some time periods; however, when emigration rates vary in time the effects of emigration should be carefully considered and accounted for, especially in comparative studies and those with policy and conservation implications.