New Insights Into the Geographical Distribution, Ecology And

Herpetology Notes, volume 14: 439-450 (2021) (published online on 26 February 2021)

New insights into the geographical distribution, ecology and conservation status of South Africa’s endemic Coastal Leaf-toed Gecko, Cryptactites peringueyi (Boulenger, 1910)

Gary K. Nicolau1,*, Melissa Petford2,3, Shelley Edwards1, Theo Busschau4, Keir Lynch5, Luke Kemp6, Jonathan Balmer1, Chad Keates1, Courtney R. Hundermark7, Joshua Weeber3,8, and Werner Conradie9,10

Abstract. Range-restricted species are generally poorly known and at higher risk of extinction than species with wider distributions. In the past, the Coastal Leaf-toed Gecko, Cryptactites peringueyi (Boulenger, 1910) caused much confusion and was once referred to as “one of the herpetological mysteries of the region”. Knowledge on the species has grown substantially, predominately due to new insights into its ecological preferences. Surveys from 2018 through early 2020 re-established the presence of C. peringueyi at three of four historical localities, as well as documenting four new localities. In 2018, preliminary data from these surveys resulted in an IUCN Red List downgrading from Critically Endangered to Near Threatened. However, with further investigation into the distribution of the species, the results from the present study more than doubled the previously estimated Extent of Occurrence from 785 km2 to 1504 km2 and the populations appear to be stable, despite environmental and anthropogenic disturbance. By incorporating environmental niche modelling, we further discuss the distribution, habits and ecology of C. peringueyi. The results presented here highlight the importance of fully understanding a species’ ecology to address its conservation status and we suggest that a new listing of Least Concern is appropriate for C. peringueyi.

Keywords. Gekkonidae, SDM, Conservation, EOO, Habitat, Ecology

Introduction Species with geographically restricted distributions are often poorly known and generally at higher risk of extinction (Söderström and Séneca, 2008; Dirnböck et 1 Department of Zoology and Entomology, Rhodes University, al., 2011; Böhm et al., 2013; Petford et al., 2019). Due Makhanda, South Africa. to increasing anthropogenic threats, it is essential to 2 South African National Biodiversity Institute, Kirstenbosch advance our knowledge of the ecological requirements Research Centre, Claremont, Cape Town, South Africa. and the full Extent of Occurrence (EOO) for range- 3 School of Animal, Plant and Environmental Sciences, restricted species and species of conservation concern University of the Witwatersrand, Johannesburg, South Africa. 4 Department of Botany and Zoology, University of (Böhning-Gaese et al., 2006; Dirze et al., 2014; Stellenbosch, Matieland, South Africa Doherty et al., 2020). As defined by the International 5 Bionerds, Environmental Consulting Services, Barrydale, Union for Conservation of Nature (IUCN), an EOO South Africa. is the area contained within the minimum continuous 6 African Snakebite Institute, Pretoria, South Africa. convex polygon drawn to encompass all known, 7 Department of Environmental Sciences, College of Agriculture inferred or projected sites of the present occurrence and Environmental Sciences, University of South Africa, Pretoria, South Africa. of a species. This metric is used to assess the measure 8 Endangered Wildlife Trust, Modderfontein, Johannesburg, of extinction risk a species may encounter across its South Africa. range (IUCN Standards and Petitions Subcommittee, 9 Port Elizabeth Museum, Beach Road, Humewood, Port 2019). Additional information regarding the ecology Elizabeth, South Africa. and accurate distribution of threatened taxa allow for 10 Department of Nature Conservation Management, Natural thorough and decisive conservation management to Resource Science and Management Cluster, Faculty of Science, George Campus, Nelson Mandela University, conserve these species and the broader ecosystems in George, South Africa. which they occur (Lamoreux et al., 2006; Söderström * Corresponding author. E-mail: [email protected] and Séneca, 2008). © 2021 by Herpetology Notes. Open Access by CC BY-NC-ND 4.0. 440 Gary K. Nicolau et al.

Cryptactites peringueyi (Boulenger, 1910) is a led to him once referring to the species as “one of the small terrestrial species of gekkonid, with a paucity herpetological mysteries in the region” (Branch, 1988). of information regarding its life history, ecology, and In 1992, 82 years after its description, C. peringueyi distribution (Branch et al., 1992; Branch and Bauer, was rediscovered 77 km west of Port Elizabeth within salt 1994; Branch, 1998; Bates et al., 2018). In addition to marsh vegetation along the banks of the lower Kromme this, the biogeography and evolutionary relationships River estuary near St. Francis Bay (Branch et al., 1992; of C. peringueyi were questioned for nearly a century Branch and Bauer, 1994). However, in early 2020, (Branch and Bauer, 1994). The species is considered a record was uploaded to the citizen science project, restricted to the naturally-fragmented Cape Seashore ReptileMap (http://vmus.adu.org.za/?vm=ReptileMAP- Vegetation (Mucina et al., 2006) in the western coastal 160104) (FitzPatrick Institute of African Ornithology, regions of the Eastern Cape Province, South Africa, 2020) of a specimen observed in 1988 at Cape St. between 0 – 30 m above sea level (Branch, 2014). Francis. This record pre-dates the original rediscovery Boulenger (1910) described this gecko based on two of the species, yet the observation was not made public specimens collected from widely separated and disjunct until recently (W. Nesser, pers. comm. 2020). In 1995, localities: a male from the ‘Little Namaqualand’ region two additional localities were documented, Willows along the west coast of South Africa (coll. Péringuey) Resort and Skoenmakerskop, with additional material in 1885 and a female, mistaken originally as a male by being collected from Chelsea Point (Branch, 1996). FitzSimons (1943), collected in 1904 (coll. Moorhouse) Between 1996 and 2009 no additional records of C. from ‘Port Elizabeth’ in the Eastern Cape Province peringueyi were documented. However, between 2010- (Branch and Bauer, 1992). Both specimens are housed 2017, several citizen science records of the species were in the South African Museum’s (SAM) collection, documented, all reported from near known localities. respectively catalogued under SAM ZR-000777 and In 1994 C. peringueyi was given a listing of SAM ZR-008628. Indeterminate (I) under then IUCN Red List categories, Re-examination of the two syntypes led to the while in 1996 the species was assigned a listing of Data designation of the Little Namaqualand specimen as the Deficient (Bates et al., 2018). Eighteen years later, C. lectotype, and by default the Port Elizabeth specimens peringueyi was reassigned to the category of Critically became the paralectotype (Branch and Bauer, 1994). Endangered (CR) during a regional threat assessment This action inadvertently implied Little Namaqualand in 2014 (Branch, 2014). This status was given as the as the type locality. This was later rectified and the Little species was considered to have a very restricted and Namaqualand locality was recorded to be ‘in error’ and severely fragmented distribution, with small estimated should be treated with caution (Branch, 1996; See Figure Area of Occurrence (AOO) and EOO, and potential 1). The paralectotype collected from Port Elizabeth was habitat threats: coastal development, fire, possible from a more likely locality, specifically Chelsea Point, coastal oil spills, and sea storms with associated a locality within the Port Elizabeth region which was flooding potentially exacerbated by climate change later found to be assigned to the specimen in the South (Branch, 2014). Shortly after the regional assessment, African Museum’s catalogue. This was a locality from the species was also assigned the status of CR in the which A. Moorhouse actively collected over his years global assessment (Branch, 2017). However, the most living in the area, thus leaving little doubt regarding the recent re-assessment of C. peringueyi assigned it the accuracy of this locality (Branch and Bauer, 1992, 1994; status of Near Threatened (NT) due to the species Branch, 1996). being documented from several additional localities, For nearly a century after the description of C. including from an official protected area, with the peringueyi, no additional material was collected only plausible threats being those from residential (Branch and Bauer, 1992) and the absence of any further and commercial development throughout the region’s observations was a cause for confusion around the status naturally fragmented habitat, temperature extremes and and origin of the species (Hewitt, 1937; FitzSimons, storm flooding (Bates et al., 2018). 1943; McLachlan, 1988; Branch and Bauer, 1992). The lack of information regarding the distribution Up until 1991, the late William R. Branch failed over and ecology of C. peringueyi motivated our aim: to a period of 12 years to re-locate this species despite gain a greater understanding of the gecko’s distribution intensive surveys in the greater Port Elizabeth region and conservation status by surveying both historical (Branch et al., 1992; Branch and Bauer, 1994). This and new localities within potentially suitable areas. New insights into South Africa’s endemic Coastal Leaf-toed Gecko 441

This was paired with the application of environmental the early mornings and late afternoons and consisted of niche modelling. We hypothesised that C. peringueyi two or more observers thoroughly searching between would occur across a much larger distributional area vegetation for signs of movement from disturbed than currently known, as suitable vegetation for this individuals, sloughed skin, or the presence of eggs. species, e.g. Cape Seashore Vegetation, appears to A variety of habitats were covered through active be more continuous along the Eastern Cape Province searching in coastal thicket vegetation, coastal dunes, coastline (Mucina et al., 2006). The increase in the salt marshes, and other coastal microhabitats between 0 estimated EOO would have further implications – 80 m elevation. for the conservation status of C. peringueyi. The identification of individuals was based on the species’ characteristic morphological features: the Material and Methods relatively large yet narrow head, a long-pointed snout with a moderately rounded yet slender body, a tapering Surveys. Between January 2018 and March 2020, 39 tail and slightly reduced adhesive pads on the digits (Fig. surveys were carried out along the coastline of the Eastern 2A; Branch, 1998). Voucher specimens were collected Cape Province of South Africa. Initial surveys focused from new and historical localities (see Appendix 1). on recording the presence of C. peringueyi at the three These specimens were humanely euthanised by injecting eastern historical localities: Chelsea Point, Willows, and with lidocaine, then fixed in 10% formalin and stored Skoenmakerskop (Fig. 1); and subsequently to search in 70% ethanol in the Port Elizabeth Museum (PEM) for new localities. Surveys took place predominantly in herpetology collection (see Appendix 1 for catalogue

Figure 1. Cryptactites peringueyi occurrence records including all known localities. Yellow circles: occurrence records from the present study; yellow square: new localities identified from the present study; red circle: citizen science and additional museum records (post-2000); purple circle: historical records (pre-2000). South Africa inset: study site (yellow circle) in relation to the general region of Little Namaqualand (turquoise circle), the locality assigned to the lectotype collected by Louis Péringuey, now widely accepted as an error. 442 Gary K. Nicolau et al.

Figure 2. (A) Adult Cryptactites peringueyi at sea level found at Skoenmakerskop. (B) Close-up of matted Syncarpha sordenscens, and an example of a communal nesting site located beneath vegetation. (C) Typical coastal habitat where C. peringueyi was recorded. Photographs taken by Gary Kyle Nicolau.

numbers). Prior to formalin fixing, liver samples were Bachman et al. (2011), and 2) by placing a minimum taken for future genetic analysis and stored in 96% convex polygon around the interpreted distribution ethanol in the PEM. Observational records from this drawn around all points and suitable habitat following study, where voucher specimens were not collected, IUCN guidelines (IUCN Standards and Petitions were uploaded to the ReptileMap database (http://vmus. Subcommittee, 2019). adu.org.za/). Occurrence points were classified according to A map of occurrence points was produced using QGIS different vegetation types in QGIS 3.4.2, according 3.4.2 (QGIS Development Team, 2019), representing to the revised and updated Vegetation Map of South all known localities, including literature and museum Africa, Lesotho and Swaziland (Dayaram et al., 2019). records, direct observation records from the present The flora that C. peringueyi appeared to be associated study, and additional citizen science records (Appendix with were identified using Goldblatt and Manning 1). Only a single record per occurrence point was (2000) and noted along with additional behavioural or included on the map, therefore excluding duplicate breeding observations of any geckos encountered. For citizen science records for known sites or inaccurate the purpose of this study and due to lack of sampling coordinates. The EOO was estimated by employing two efforts between known or newly documented localities, different methods: 1) by incorporating all the compiled we could not assign material to separate populations, point localities occurrences (Appendix 1, excluding thus occurrence records within 20 km of each other the Little Namaqualand locality) in the open-source have been assigned to the same range, with different Geospatial Conservation Assessment Tool (GeoCAT ranges separated by > 20 km. - http://geocat.kew.org), following the methods in New insights into South Africa’s endemic Coastal Leaf-toed Gecko 443

Environmental Niche Modelling. The potential The environmental niche model was constructed using suitable range of C. peringueyi was predicted using an the presence-background Maxent (Phillips et al., 2006) environmental niche model, conducted along a 450 km technique due to its credibility in modelling species with stretch of coastline between Wilderness (Western Cape small sample sizes and presence-only records (Elith et Province) in the west, to Begha (Eastern Cape Province) al., 2006, 2010; Pearson et al., 2007). As parameter in the east. The extent of the study area was selected by settings can have a considerable influence on model calculating the largest distance between the most western outcomes, species-specific tuning was conducted using and eastern records and extending the study area with the ENMevaluate function of the ENMeval R package the calculated distance (160 km) on either side, with to identify the best model for C. peringueyi (Anderson the known localities of C. peringueyi at the centre. This and Gonzalez, 2011; Muscarella et al., 2014). Models range was selected as it encompasses the current known were constructed with different levels of regularisation range of C. peringueyi and allows for predictions of (0.5 to 4.5 in 0.5 increments) and combinations of the the potential distribution within a reasonable distance feature classes (linear (L); quadratic (Q); hinge (H); of known localities, considering the species is range- product (P) and threshold (T)). As there were fewer restricted and without prior knowledge on dispersal than 25 occurrence points, the data were partitioned ability, biotic interactions or potential barriers. All into testing and training bins using the “jackknife” analyses were conducted in R Studio version 3.5.1 (R method. To account for any spatial sampling bias in the Core Team, 2018) unless stated otherwise. To reduce study area, 10 000 background points were randomly spatial autocorrelation, the species occurrence points selected from a bias raster, which was constructed with were spatially rarefied using the thinning function of a dataset of 12 348 records of all reptile species from the R package spThin (Aiello-Lammens et al., 2015) to within the bounding box of the modelling study area. a distance of 1 km, with 16 records retained for further These bias records were thus compiled from the Global use in the modelling. Biodiversity Information Facility (GBIF.org, accessed 5 Nineteen bioclimatic variables (www.worldclim.org) April 2020). Although GBIF datasets can contain errors, and elevation (GTOPO30) (Earth Resources Observation these records were not explicitly used in the actual and Science Center, 1997) were downloaded at a 30 modelling for C. peringueyi, but rather used to create arc-second resolution, while vegetation layers at a scale a bias layer to account for potential spatial bias within of 1:1 000 000 (South African National Biodiversity the study area. Therefore, it was not as important that Institute, 2018) were downloaded. The vegetation layer all of the records gathered from the GBIF platform have was categorised into 49 broader vegetation types. A the same accuracy level than those used for the study Pearson’s correlation coefficient test was performed species as only general trends were needed to account on the bioclimatic variables using the cor.test function for spatial sampling bias. of the R stats package (R Core Team, 2018) to reduce The optimum model for C. peringueyi was selected by collinearity, as strongly correlated variables make it evaluating a variety of model metrics. Firstly, to select difficult to interpret the importance of predictor variables a model with a high goodness of fit, yet not overly (Dormann et al., 2013). Variable pairs with a correlation complex, the model with the lowest Akaike Information coefficient ≥ 0.7 were inspected. Only one variable from Criterion (AIC), corrected for small sample sizes, was each correlated pair were retained based on general selected (Burnham and Anderson, 2004). Following biological requirements of lizards and primary limiting this, to ensure that the model with the lowest AIC factors for ectotherm distributions (Araújo et al., 2006; was not overfitting, the minimum training omission

Vitt and Caldwell, 2013; Pintor et al., 2016; Kearney et rate (ORmtp), the training omission rate (OR10), the al., 2018). We specifically prioritised temperature and threshold-independent metric (AUC) and the difference precipitation variables and those that relate to extremes between test and training AUC (AUCdiff) were inspected and variability (Araújo et al., 2006; Bradie and (Anderson and Gonzalez, 2011). Finally, the resulting Leung, 2016). The variables which were included for map and response curves were inspected for ecological analysis were: mean diurnal temperature range (Bio2); realism. Once the optimal model was selected, the isothermality (Bio3); mean temperature of the wettest permutation importance of variables was inspected quarter (Bio8); mean temperature of the coldest quarter to identify the most important variables for the (Bio11); annual precipitation (Bio12); precipitation environmental niche model. To investigate the potential seasonality (Bio15), precipitation of the coldest quarter range size of C. peringueyi, the distribution map was (Bio19) and vegetation type. converted into a binary suitable/unsuitable map using 444 Gary K. Nicolau et al. the 10-percentile training value (Liu et al., 2005). The records documented in the present study Following this, the potential extent of the distribution are distributed across four broad vegetation types, was estimated in QGIS 3.4.2. using the GRASS r.report including the Albany Alluvial Vegetation, Cape function. Seashore Vegetation, Elands Forest Thicket, St. Francis Dune Thicket, and on the edges of non-terrestrial Results estuarine functional zones (Appendix 1). Whilst C. peringueyi was predominantly found within or under Surveys. The records included in the present study plants of the species Syncarpha sordenscens, that form make up ten broad localities and consist of 42 records, a mat-like covering (Fig. 2B–C), individuals were also including 14 historical literature and museum records, recorded within Phragmites sp. and Juncus sp. clumps 18 direct records from the present study, and ten citizen science records from the ADU ReptileMap (Fig. 1, along the Kromme River estuary and under a variety of Appendix 1). These localities occur in three broad debris, both natural and artificial, in the urban localities areas: 1) an eastern range comprising Kini Bay, Sardinia of the central range. Bay, Skoenmakerskop, Willows, and Chelsea Point; Throughout the surveys, multiple pairs of hatched, 2) a central range comprising Oyster Bay, Cape St. freshly laid or developing eggs were observed clustered Francis, St. Francis Bay, and Kromme River estuary; together between and within matted coastal vegetation, and 3) a new western range in Eersterivierstrand. A like that of S. sordenscens (Fig. 2B), while other record located within the Seal Point Nature Reserve observations of nesting sites were made within the in the Cape St. Francis locality represents the first and vegetation of urban gardens. It was also noted that the only confirmed occurrence within an official protected species displayed crepuscular, nocturnal, and diurnal area, although the interpreted distribution incorporates activity. additional protected areas. Regardless of numerous Environmental Niche Modelling. When the attempts, C. peringueyi was not observed at the environmental niche models were run including the historical Chelsea Point locality. The latest unsuccessful categorical variable vegetation, the evaluation metrics surveys at Chelsea Point were conducted during March of the model with the lowest AIC value indicated 2020. The Willows and Skoenmakerskop localities, 2.5 overfitting, with high ORmtp (0.13) and OR10 (0.19). km east of Chelsea Point, appeared to have abundant Further inspection of the resulting distribution map C. peringueyi, with direct searches revealing up to 20 confirmed that the model, including vegetation, was individuals. overfit, with poor model accuracy around occurrence The surveys resulted in four new localities: Sardinia points, many of which were excluded from potentially Bay, Kini Bay, Oyster Bay and Eersterivierstrand. suitable areas. Following this, the models were rerun The Kini Bay and Oyster Bay localities extend the excluding vegetation type and thus included only distribution in the eastern range by 14.5 km west and bioclimatic variables. The removal of vegetation type the central range by 16 km west respectively. The record from our analyses is justified due to its poor model from Eersterivierstrand represents a new westerly performance and the fact that our surveys indicate that locality, ± 40 km west of the new Oyster Bay records. C. peringueyi occurs across four broad vegetation types, With the inclusion of the additional three new localities, some of which also occur in areas far outside of our the eastern and central ranges occupy ± 25 km and ± 30 predicted distribution model (Fig. 3). The bioclimatic- km of the Eastern Cape Province coastline, respectively, only model with the lowest AIC value appeared to and up to 7 km inland in the Kromme River estuary. perform well, with high AUC (0.98) and low ORmtp

The species’ range, now extends over five quarter- (0.06), OR10 (0.13) and AUCdiff (0.01) indicating that degree grid cells (QDGC, see Bates et al., 2014 on the resulting model was not overfitted to the occurrence the use of QDGC); 3425BA (Sardinia Bay, Willows, points. Inspection of the distribution map indicated Skoenmakerskop and Chelsea Point), 3424BB (Cape a good fit in terms of known occurrence points. As a St. Francis, St. Francis Bay and Kromme River estuary), result of the good evaluation metrics and apparent 3425AB (Kini Bay), 3424AA (Eersterivierstrand) and model fit, the bioclimatic-only model was retained for 3424BA (Oyster Bay) (Fig. 1). The updated EOO further analyses. estimated in GeoCAT using point locality occurrences The predicted distribution of C. peringueyi was was 1243 km2, while the EOO using the interpreted restricted to the coastal areas (Fig. 3A). In addition distribution was estimated at 1504 km2. to the known areas of occurrence, the model suggests New insights into South Africa’s endemic Coastal Leaf-toed Gecko 445

Figure 3. (A) Suitability map showing the potential extent of the distribution of Cryptactites peringueyi for the optimum Maxent model as determined by the program ENMeval in R. Predicted suitability is displayed on a scale of low (blue) to high (red). (B) Binary suitable/unsuitable map showing the suitable range of Cryptactites peringueyi. The binary map was created using the optimum Maxent model as derived from the package ENMeval in R and implementing a 10-percentile training value (Liu et al., 2005). Grey – unsuitable; Green – suitable; Pink Circles – occurrence records of C. peringueyi; Yellow Circles – major towns.

that there are suitable areas for C. peringueyi to the predicted distribution is positively correlated with the west in the Sedgefield area and in two areas eastwards, three highest contributing variables, mean temperature between Colchester and Port Alfred. These three new of the coldest quarter (Bio11), precipitation of the areas, as highlighted by the niche model (Fig. 3B), are coldest quarter (Bio19) and annual precipitation (Bio12), disjunct from the current known distribution by ± 115 predicting higher suitability in areas with warmer mean km west for the former, and ± 40 km and ± 100 km east, temperatures and higher rainfall during the coldest three respectively, for the latter. The binary presence/absence months. map suggests that the total predicted area or potential extent of the distribution within the study area is 1035 Discussion km2 (Fig. 3B). The predicted suitable area within the known distribution of the species, between Chelsea Through the use of the environmental niche modelling Point and Eersterivierstrand, is only 755 km2. and our surveys, we have significantly increased the The main environmental variable contributing to the knowledge regarding the geographical distribution and predicted distribution of C. peringueyi, as indicated by conservation status of Cryptactites peringueyi. The re- the permutation importance of variable percentages, confirmation of the gecko at three of the four historical was mean temperature of the coldest quarter (62.80%) sites and the addition of four new localities were the followed by precipitation of the coldest quarter (10.85%), key findings, indicating that the species is distributed annual precipitation (8,90%), mean temperature of over a larger range than previously documented. The the wettest quarter (5.78%), precipitation seasonality inclusion of the new localities and additional occurrence (4.22%), mean diurnal temperature range (1.10%), and points more than doubled the previous EOO, with mean temperature of the wettest quarter (<0.01%). The estimations of 1243 km2 (point localities) and 1504 km2 446 Gary K. Nicolau et al.

(interpreted distribution). The EOO using the interpreted No. 24 of 2008), and thus further urban development distribution is likely more accurate than the estimated should not have any severe impact on the species. EOO using point locality occurrences, which is likely The risk of potential flooding due to severe climatic underestimated when taking into account the results events (Branch and Bauer, 1994) is unlikely, as specimens of the environmental niche model. Together, these have been recorded up to 7 km inland along the Kromme observations support the conclusion that the gecko is River estuary and away from rivers. Although historical not as restricted nor as threatened as previously stated. surveys after a major flooding event at the Kromme The most recent IUCN Red List re-assessment of C. River locality were unsuccessful (Haagner et al., peringueyi used unpublished results and downgraded the 1996), contemporary records with multiple occurrence species from Critically Endangered to Near Threatened points at this locality allude to the species’ resilience (Bates et al., 2018). The re-assessment incorporated our to sporadic flooding (Fig. 1). The impact of climatic new records from urban areas in Cape St. Francis and change is further mitigated by the occurrence of C. St. Francis Bay, and at Willows and Skoenmakerskop, peringueyi across four broad vegetation types and on the first observations of C. peringueyi from the eastern the edges of non-terrestrial estuarine functional zones, range in 23 years (Bates et al., 2018). Our two new indicating a degree of tolerance to vegetation type localities at Kini Bay and Oyster Bay and the occurrence and that the species is less habitat specific as defined of the species in an official protected area at Seal Point by vegetation type than previously thought. However, Local Authority Nature Reserve were also included although the species occurs across four broad vegetation (Bates et al., 2018). However, shortly after the re- types which cover large areas (Dayaram et al., 2019), assessment was published, we identified a new locality the correlative niche model suggests that the species near the intertidal zone of the Marine Protected Area at is still primarily restricted to the coastal regions (Fig. Sardinia Bay and the major westerly range-extension 3). The largest contributing variable to the niche model from Eersterivierstrand. was the mean temperature of the coldest quarter, with Inferred threats to C. peringueyi included coastal warmer areas being more suitable. This is expected as developments, with associated habitat loss such as ectothermic organisms have a strong reliance on external through housing development and urbanisation, and environmental conditions to regulate body temperatures climatic events, such as flooding and global climate (Avery, 1982). Species’ ranges that are highly influenced change (Branch, 2014, 2017; Bates et al., 2018). by temperature are also likely to be more restricted in However, several observations have shown that the range, due to their physiological requirements (Aragón species displays a level of tolerance to anthropogenic et al., 2010). disturbance. Individuals have recently been observed in In addition to the known localities, the correlative the urban areas of St. Francis Bay and Cape St. Francis, niche model predicted the range to be far greater than both during this study and through citizen science what is currently known and not as fragmented (Fig. observations, climbing on walls and breeding amongst 3B). The model is, however, predictive and represents garden vegetation. Historical records provide further the potential extent of the distribution; therefore, C. evidence of the species’ tolerance to anthropogenic peringueyi may or may not occur in the areas predicted disturbance over time. The first records of the species to be suitable. Although the model had a high AUC at the Skoenmakerskop locality specified suburban value suggesting good model fit, this high AUC is likely gardens (Branch, 1996), and several clutches of due to C. peringueyi’s restricted range in comparison to eggs were discovered under a 200-litre drum at the the modelled study area (Phillips, 2006). Despite these Kromme River estuary locality (Haagner et al., 1996). points, the model does provide a useful guideline to These observations are important when considering a investigate the distribution of C. peringueyi. restricted species’ response to disturbance (Doherty et Preliminary surveys were conducted in several areas al., 2020), particularly since the species is still present highlighted as potentially suitable, including Cannon at these localities at the time of writing. In addition to Rocks, Kenton on Sea, Port Alfred, Seaview (2.5 this, the Integrated Coastal Management Act of 2008 on km west of Kini Bay), Van Stadens River Mouth, coastal development states one may not develop within Gamtoos River Mouth, Tsitsikamma, Seekoeivlei (6 km 1 km of the high-water mark for non-zoned coastal south of Jeffreys Bay) and Robberg (Fig. 3B). These development areas, and within 100 m of previously non-intensive preliminary surveys were, however, zoned development areas (National Environmental unsuccessful and future surveys would be needed to Management: Integrated Coastal Management Act, Act confirm the presence of the species along the outlying New insights into South Africa’s endemic Coastal Leaf-toed Gecko 447 coastal regions of the model. The model also highlighted localities. Although our results have increased our the area surrounding Sedgefield as potentially suitable. knowledge of the distribution of C. peringueyi, it is Sedgefield is 115 km away from the nearest recorded evident that further research is still required, especially locality, separated by a large area of low probability of regarding population size, demography, dispersal, and occurrence and the maximum distance between known habitat preference. We hope this work may act as a localities is roughly 40 km, leading us to question the catalyst for future research into this unique species. likelihood of C. peringueyi occurring there (Fig. 3B). However, palaeoceanographic models suggest that Acknowledgements. We would like to thank the Department historically these coastal regions, and thus suitable of Economic Development, Environmental Affairs and Tourism habitat, may have had a far greater and less fragmented for permits (permit numbers: CRO 43/17CR and CRO 44/17CR, range (Grobler et al., 2020). Therefore, the occurrence CRO 134/19CR and CRO135/19CR) issued to WC. Ethical of C. peringueyi around Sedgefield is plausible and clearance for this study was granted from the Port Elizabeth Museum (Ethical Clearance no. 2017-2). Authors would further further investigation would be needed to confirm the like to thank Emily Anne Jackson, Justin Rhys Nicolau, Dewald presence or absence of the species. Swanepoel, Ryan van Huyssteen, Luke and Ursula Verburgt, It is important to note that the species appears to be Francois Theart, Javier Lobon Rovira and Walter Neser who breeding in a variety of different habitats. This is evident either assisted in surveys or supplied additional records during from the number of communal nesting sites discovered this study. We thank the two reviewers, Krystal Tolley and Aaron during the present study. Communal breeding has Bauer, whose inputs helped improve the quality of the article. historically been recorded for C. peringueyi in a dense clump of grass at the Kromme River estuary locality References and a single clutch was also discovered under a log Aiello-Lammens, M.E., Boria, R.A., Radosavljevic, A., Vilela, in the Sarcocornia perennis floodplains of the estuary B., Anderson, R.P. (2015): spThin: an R package for spatial (Haagner et al., 1996). Throughout the study, communal thinning of species occurrence records for use in ecological breeding sites were primarily observed amongst niche models. Ecography 38: 541–545. Syncarpha sordenscens at the coastal localities and Anderson, R.P., Gonzalez, J.I. (2011): Species-specific tuning within garden vegetation or under debris in urban areas. increases robustness to sampling bias in models of species These breeding observations are further indications distributions: an implementation with Maxent. Ecological of the species’ adaptability to different habitats and Modelling 222(15): 2796–2811. resilience to disturbance. It is, therefore, likely that the Aragón, P., Lobo, J.M., Olalla-Tárraga, M.Á., Rodríguez, M.Á. (2010): The contribution of contemporary climate to ectothermic species may be utilising a greater variety of habitats and endothermic vertebrate distributions in a glacial refuge. than currently recognised and that existing observations Global Ecology and Biogeography 19: 40–49. are biased to habitats that are easy to survey. Araújo, M.B., Thuiller, W., Pearson, R.G. (2006). Climate warming Our findings suggest that C. peringueyi should and the decline of amphibians and reptiles in Europe. 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Appendix 1. Voucher specimens and records included in the present study.

Voucher Latitude Longitude QDGC Date Location Vegetation Type Collectors and observers

Lectotype NA NA NA 1885 Little Namaqualand “In Error” NA Péringuey, L.

Paralectotype -34.03 25.38 3425BA 02.12.1904 Chelsea Point, Port Elizabeth St. Francis Dune Thicket Moorhouse, A.

ReptileMAP: 160104 -34.2087 24.8266 3424BB 05.01.1988 Render Lane, Cape St. Francis St. Francis Dune Thicket Neser, W; Kritzinger, J.

PEM R6886 -34.1166 24.7833 3424BB 03.02.1992 Kaia de Balaia, upper Kromme River Elands Forest Thicket Craig, G. estuary, Humansdorp

PEM R11334 -34.1361 24.8027 3424BB 29.03.1992 Kromme River Bridge, Humansdorp Estuarine Functional Zone Burger, M.

PEM R7210 -34.1166 24.7833 3424BB 26.07.1992 Kaia de Balaia, upper Kromme River Elands Forest Thicket Craig, G. estuary, Humansdorp

PEM R6908 -34.1416 24.8061 3424BB 11.01.1992 200m west of Kromme River bridge, Estuarine Functional Zone Hall, R. Humansdorp

PEM R6912 -34.1333 24.8000 3424BB 11.01.1992 50m northwest of Kromme River bridge, Estuarine Functional Zone Branch, W; Ralfe, M; Hall, R. Humansdorp

PEM R7210 -34.1166 24.7833 3424BB 26.07.1992 Kaia de Balaia, upper Kromme River Elands Forest Thicket Craig, G. estuary, Humansdorp

PEM R8068 -34.1500 24.8166 3425BB 06.02.1993 Kromme River Mouth, western bank, St. Francis Dune Thicket Branch, W. Humansdorp

PEM R10875 -34.1402 24.8111 3424BB 07.01.1995 Bridge at Kromme River Mouth, Estuarine Functional Zone Haagner, G.V. Humansdorp

PEM R12206 -34.0447 25.6325 3425BA 17.07.1995 Chelsea Point, Port Elizabeth St. Francis Dune Thicket Riley, D.

PEM R12207 -34.0452 25.6183 3425BA 17.07.1995 Willows Resort, Port Elizabeth St. Francis Dune Thicket Riley, D.

PEM R12209 -34.0413 25.5360 3425BA 11.05.1995 Skoenmakerskop, Port Elizabeth St. Francis Dune Thicket Riley, D.

ReptileMAP: 8665 -34.1908 24.8368 3424BB 25.11.2010 St. Francis Airpark, Cape St. Francis St. Francis Dune Thicket Darling, G.

PEM R19186 -34.1377 24.8077 3424BB 02.02.2011 West of the bridge crossing Kromme River, Estuarine Functional Zone Weldon, C; Conradie, W; Botha, St. Francis Bay V.

ReptileMAP: 153218 -34.1588 24.8113 3424BB 10.11.2014 Erf 178, St. Francis Links, St. Francis Bay St. Francis Dune Thicket Logie, B; Logie, C.

ReptileMAP: 153392 -34.1675 24.8297 3424BB 09.02.2015 7 Edward Rd, St. Francis Bay St. Francis Dune Thicket Logie, B; Logie, C.

ReptileMAP: 153395 -34.1612 24.8177 3424BB 16.02.2015 8 St. Francis Links, St. Francis Bay St. Francis Dune Thicket Logie, B; Logie, C. 450 Gary K. Nicolau et al.

Appendix 1. Continued.

Voucher Latitude Longitude QDGC Date Location Vegetation Type Collectors and observers ReptileMAP: 153529 -34.1733 24.8368 3424BB 26.01.2015 6 Juan Carlos Crescent, Santareme, St. St. Francis Dune Thicket Logie, B; Clause, L. Francis Bay

ReptileMAP: 155758 -34.1908 24.8368 3424BB 26.11.2015 St. Francis Airpark, Cape St. Francis St. Francis Dune Thicket Darling, G.

ReptileMAP: 159823 -34.1908 24.8368 3424BB 16.11.2016 St. Francis Airpark, Cape St. Francis St. Francis Dune Thicket Darling, G; Darling, D.

ReptileMAP: 160176 -34.1908 24.8368 3424BB 13.12.2016 St. Francis Airpark, Cape St. Francis St. Francis Dune Thicket Darling G.

ReptileMAP: 166990 -34.1863 24.8227 3424BB 28.11.2017 Cape St. Francis dump St. Francis Dune Thicket Hundermark, C; Kemp, L; Keates, C.

PEM R24283 -34.0452 25.6183 3425BA 28.01.2018 East of Willows Resort, Port Elizabeth St. Francis Dune Thicket Present study: Nicolau, G.K; Kemp, L.

PEM R24285 -34.0455 25.6185 3425BA 29.01.2018 East of Willows Resort, Port Elizabeth St. Francis Dune Thicket Present study: Nicolau, G.K.

ReptileMAP: 165044 -34.0455 25.6185 3425BA 08.02.2018 East of Willows Resort, Port Elizabeth St. Francis Dune Thicket Present study: Kemp, L; Nicolau, G.K.

ReptileMAP: 174435 -34.2108 24.8307 3424BB 22.06.2018 Seal Point Nature Reserve, Cape St. Francis St. Francis Dune Thicket Present study: Lynch, K.

PEM R24284 -34.0304 25.3959 3425AB 01.07.2018 1 km east of Kini Bay St. Francis Dune Thicket Present study: Conradie, W; Hundermark, C.

PEM R24295 -34.1763 24.6639 3424BA 15.08.2018 Oysterbay, 16 km west of Cape St. Francis St. Francis Dune Thicket Present study: Conradie, W; Keates, C; Busschau, T; Edwards, S; Weeber, J.

PEM R24294 -34.0345 25.5050 3425BA 25.08.2018 Sardinia Bay, Port Elizabeth St. Francis Dune Thicket Present study: Nicolau, G.K.

ReptileMAP: 174427 -34.0345 25.5050 3425BA 25.08.2018 Sardinia Bay, Port Elizabeth St. Francis Dune Thicket Present study: Nicolau, G.K.

ReptileMAP: 174428 -34.0413 25.5360 3425BA 25.08.2018 Skoenmakerskop, Port Elizabeth St. Francis Dune Thicket Present study: Nicolau, G.K.

ReptileMAP: 174431 -34.0413 25.5360 3425BA 28.09.2018 Skoenmakerskop, Port Elizabeth St. Francis Dune Thicket Present study: Nicolau, G.K; Jackson, E.A; Nicolau, J.R; Swanepoel, D.

PEM R24979 -34.0719 24.2183 3424AA 29.11.2018 Eersterivierstrand, 33 km Southeast of Cape Seashore Vegetation Present study: Balmer, J. Stormsriver

PEM R25010 -34.0452 25.6194 3425BA 01.02.2018 East of Willows Resort, Port Elizabeth St. Francis Dune Thicket Present study: Nicolau, G.K.

PEM R25071 -34.0450 25.6147 3425BA 16.01.2019 East of Willows Resort, Port Elizabeth St. Francis Dune Thicket Present study: Conradie, W; Rovira, J.L.

ReptileMAP: 174425 -34.0450 25.6147 3425BA 09.08.2018 East of Willows Resort, Port Elizabeth St. Francis Dune Thicket Present study: Nicolau, G.K.

ReptileMAP: 174429 -34.0413 25.5360 3425BA 11.05.2018 Skoenmakerskop, Port Elizabeth St. Francis Dune Thicket Present study: Nicolau, G.K.

ReptileMAP: 175319 -34.1994 24.8310 3424BB 12.09.2019 Next to Full Stop Café, Cape St. Francis St. Francis Dune Thicket Present study: Kemp, L; Keates, C; Theart, F; Petford, M; Van Huysteen, R.

ReptileMAP: 175748 -34.1983 24.8363 3424BB 12.09.2019 Cape St. Francis resorts, Cape St. Francis St. Francis Dune Thicket Present study: Verburgt, L; Verburgt, U; Conradie, W.

ReptileMAP: 174426 -34.0450 25.6147 3425BA 19.03.2020 East of Willows Resort, Port Elizabeth St. Francis Dune Thicket Present study: Nicolau, G.K.

Accepted by Graham Walters