Published in the United States of America 2023 * VOLUME 17 * NUMBER 1/2 AMPHIBIAN & REPTILE ONSERVATION amphibian-reptile-conservation.org ISSN: 1083-446X eISSN: 1525-9153 Front cover: Falk’s Blue-headed Tree Agama (Acanthocercus cf. cvanocephalus) from south-eastern Angola. Photographed during an expedition to elucidate the biodiversity of south eastern Angola with the Okavango Wilderness Project, in association with National Geographic. A different photograph of this individual appeared as Fig. 2 in an article cataloging the lizards identified during that Project, which was published as: Amphibian & Reptile Conservation 16(2) [General Section]: 181—214 (e322). Photo by Chad Keates. Official journal website: amphibian-reptile-conservation.org Amphibian & Reptile Conservation 17(1/2) [General Section]: 1-18 (e324). An annotated checklist of the herpetofauna of the Sibiloi National Park in northern Kenya based on field surveys 1.2,Sebastian Kirchhof, *Victor Wasonga, “Tomas Mazuch, *Stephen Spawis, and * Patrick Kinyatta Malonza ‘Division of Science, New York University Abu Dhabi, Abu Dhabi, UNITED ARAB EMIRATES ?Museum fiir Naturkunde, Leibniz Institute for Evolution and Biodiversity Science, Invalidenstr. 43, 10115 Berlin, GERMANY +Herpetology Section, National Museums of Kenya, P.O. Box 40658-00100, Nairobi, KENYA *Department of Forest Ecology, Mendel University in Brno, 61300, Brno, CZECH REPUBLIC °%7 Crostwick Lane, Spixworth, Norwich, NRIO 3PE, UNITED KINGDOM Abstract.—The xeric Lake Turkana area in northern Kenya is often referred to as the “Cradle of Mankind” due to the abundance of hominin fossils. Sibiloi National Park in the Turkana Basin has been well studied for its fossils, but the extant biodiversity of the park remains largely under-surveyed. Today, the ecosystem is threatened by climate change, increasing human population pressure, poaching pressure, overgrazing by domestic stock, and a series of major hydropower dams and irrigated agricultural schemes (particularly the Gilgel Gibe Ill Dam) in Ethiopia, which may have a negative impact on the water supplies of the lake. The Turkana Basin has a high diversity of arid land herpetofauna, particularly terrestrial geckos. However, due the region’s remoteness many expected species have never been recorded within the National Park. Here we provide an annotated list of the rich reptile and amphibian fauna based on two recent field surveys, including multiple first records for Sibiloi National Park. The surveys yielded records for 34 species, including six amphibians and 28 reptiles (one fresh water turtle, one crocodile, 18 lizards, and eight snakes). In total, 49 species of herpetofauna are currently known for Sibiloi National Park, including eight amphibians and 41 reptiles (three fresh water turtles, one crocodile, 25 lizards, and 12 snakes). Of those, five species are protected by the Convention on International Trade in Endangered Species (CITES; Appendices I, Il, or Ill), i-e., Trionyx triunguis, Crocodylus niloticus, Varanus albigularis, Varanus niloticus, and Eryx colubrinus (all Appendix Il). Three species are listed on the International Union for Conservation of Nature (IUCN) Red List (Vulnerable, Data Deficient), i.e., Sclerophrys turkanae, Trionyx triunguis, and Pelusios broadleyi. Two species (one toad, Sclerophrys turkanae, and one terrapin, Pelusios broadleyi) are endemic to Kenya and most likely endemic to the vicinity of Lake Turkana. Overall, the herpetofauna of the Sibiloi National Park already seems to be negatively affected and is further threatened by climate change and land use activity. Furthermore, the area comprises a number of CITES listed and IUCN Red List species other than the herpetofauna, including endemics, that warrant protection and conservation measures to prevent further defaunation. Keywords. Amphibians, climate change, conservation, field body temperature, Lake Turkana, land use, pastoralism, reptiles, Turkana Basin Institute Citation: Kirchhof S, Wasonga V, Mazuch T, Spawis S, Malonza KP. 2023. An annotated checklist of the herpetofauna of the Sibiloi National Park in northern Kenya based on field surveys. Amphibian & Reptile Conservation 17(1/2) [General Section]: 1-18 (e324). Copyright: © 2023 Kirchhof et al. This is an open access article distributed under the terms of the Creative Commons Attribution License [Attribution 4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org. Accepted: 3 November 2022; Published: 1 June 2023 changes, pollution, and climate change. Such threats are linked to both anthropogenic activities and natural causes. In East Africa, the distribution ranges of Introduction Reptiles and amphibians are currently considered to be among the world’s most endangered groups, with 40.7% of amphibians and 21.1% of reptiles threatened with extinction (Cox et al. 2022). This is perhaps attributed to the fact that they are sensitive to habitat destruction and fragmentation, various environmental Correspondence. sebkirchhof@gmail.com, malonzapk@gmail.com Amphib. Reptile Conserv. many amphibians and reptiles are still poorly known due to the vast areas that have yet to be explored by herpetologists (Spawls et al. 2018; Tolley et al. 2016). Nevertheless, even the limited herpetological research conducted to date clearly indicates a high diversity of amphibians and reptiles in East Africa, including June 2023 | Volume 17 | Number 1/2 | e324 The Herpetofauna of Sibiloi National Park, Kenya many threatened and endemic species (Channing and Howell 2006; Largen and Spawls 2010; Spawls et al. 2018, 2019; Tolley et al. 2016). Sibiloi National Park (SNP) is located in Marsabit County in northern Kenya and about 1,570 km? in size (Fig. 1). The park lies in the xeric Gregory Rift (the eastern branch of the East African Rift fracture system) on the northeastern shore of Lake Turkana (formerly Lake Rudolf), the largest permanent desert lake and the largest alkaline lake on Earth. The lake is part of an endorheic basin, with three rivers (Omo, Turkwel, and Kerio) supplying most of the lake’s water, and it has no outflow. The area is often referred to as the “Cradle of Mankind” because it is quite famous for its hominin fossils, in particular Australopithecus and early Homo fossils. The park was established in 1973 by the government of Kenya, mainly for the protection of its paleontological sites, and was named for Mount Sibiloi which is located at its southern boundary. Together with the South Island and the Central Island National Parks, the Lake Turkana National Parks were declared as a UNESCO World Heritage site in 1997. The site was listed as a World Heritage Site in Danger in 2018, mostly owing to large-scale transfrontier hydrological projects, in addition to climate change, increasing human population pressure, poaching pressure, and overgrazing by domestic stock. The Lake Turkana area is characterized as a desert and xeric shrubland biome (Somali-Maasai xeric grasslands and shrublands ecoregion; Olson et al. 2001), and contains a variety of different habitats (Fig. 2). The area is surrounded by Acacia (now Vachellia)- Commiphora bushland and thicket (Olson et al. 2001), bordering the Chalbi Desert to the East, and elevations range from around 340 m to 550 m (Thorsell 2003). The plains are dominated by sand, silt, and gravel and are interspersed with volcanic formations, which include Mount Sibiloi and its remains of an approximately 7 MYA old petrified forest (Thorsell 2003). The climate is generally hot and dry, with the annual rainfall of about 250 mm mainly restricted to March and April (Thorsell 2003). Temperatures at the nearest weather station in Lodwar on the western side of Lake Turkana average 32 °C throughout the year, with an average annual maximum of 37 °C and average annual minimum of 26 °C (Mbaluka and Brown 2016). There are six major ethnic groups living adjacent to the eastern side of the lake, including the Daasanach, Gabbra, Turkana, Elmolo, Rendille, and the Samburu (Kaijage and Nyagah 2009). Pastoralism (mainly goats and sheep, but also cattle, donkeys, and camel), agro-pastoralism, and fishing are the main sources of livelihood for the local population. Due to the ever-growing population around the lake, these land use practices together with frequently occurring droughts have a degrading impact on the habitats of the area. The vegetation is dominated by perennial and annual grassy plains and dwarf shrublands, covered with the grass Aristida sp. and the common dwarf shrubs Indigofera spinosa and Duosperma longicalyx (Mbaluka and Brown 2016). Much of the lake Shoreline is occupied by perennial grasslands with the halophyte Sporobolus spicatus and Dactyloctenium sp. (Mbaluka and Brown 2016). However, large parts of the shoreline in the study area are completely devoid of vegetation or scattered with dead reed tussocks in some areas. Woodlands and forests are only associated with ephemeral streams and are dominated by Vachellia tortilis (Mbaluka and Brown 2016). Legend © Sampling sites [_] Country borders oo Sibiloi National Park boundaries WWF terrestrial ecoregions [GS Masai xeric grasslands and shrublands Gl Ethiopian montane grasslands and woodlands GB East African montane forests [4 Ethiopian montane forests HS Northern Acacia-Commiphora bushlands and thickets [9 Somali Acacia-Commiphora bushlands and thickets (0) Lake Turkana Fig. 1. Location of Sibiloi National Park (UNEP-WCMC and IUCN 2022) in Kenya and the main study sites: IL (IIkemere), KA (Karare), KF (Koobi Fora), LO (Lomosia), AB (Alia Bay), and TBI (Turkana Research Institute). The inset map shows the African continent, and the black square indicates the location of the enlarged map. Amphib. Reptile Conserv. 2 June 2023 | Volume 17 | Number 1/2 | e324 Kirchhof et al. According to the latest IUCN Conservation Outlook Assessment for the area (IUCN World Heritage Outlook 2020), the local mammal fauna is largely depleted. Notably, the hippopotamus, wild dog, lion, cheetah, Reticulated Giraffe, Grevy’s Zebra, and Lelwel Hartebeest are species which were formerly abundant there but are now considered rare, endangered, or locally extinct. Historical data indicate that the lake once supported the world’s largest colony of Nile Crocodile, but recent field observations suggest the crocodile population is a fraction its former size, likely due to increasing anthropogenic pressures (IUCN World Heritage Outlook 2020). The lake is rich in fish (47 species; Birdlife International 2022). In addition, Lake Turkana is an internationally recognized Important Bird Area (Birdlife International 2022), with 84 water bird species, including 34 Palaearctic migrants, according to the latest available data. The only known extensive reptile and amphibian survey of the area was conducted by Ziliani et al. (2006). The results were presented at the 6" Congresso Nazionale della Societas Herpetologica Italica in 2006, but have not been formally published other than the abstract of that presentation and a new species description (Sindaco et al. 2007). The annotated checklist provided here is the result of two six-week herpetological surveys of the SNP conducted in 2016 and 2017. In addition, the species recorded by Ziliani et al. (2006) are included and discussed. The identifications of the new materials collected in the present study were based on the species descriptions and other literature, morphological comparisons using the reference collection at the National Museums of Kenya (NMK) and, in some cases, the advertisement calls of amphibians (S. Kirchhof, unpub. data). Material and Methods Two expeditions to SNP were conducted from 7 November 2016 to 11 December 2016 and from 28 March 2017 to 24 April 2017. Three observers sampled a total of five sites across the National Park, three along the Lake Turkana (L) shoreline, 1.e., Alia Bay (AB), Koobi Fora (KF), and Ilkemere (IL), and two further inland, 1.e., Karare (KA) and Lomosia (LO) (Fig. 1). The LO site was only surveyed in March/April 2017. At each site close to the lake, we walked three transects (one 1n grassland (G) along the shore, one in bushland (B), and one in a dry riverbed (R); Fig. 2), each with a length of about 800—1,000 m, for 3 hours in the morning from 0800-1100 h and 4 hours in the evening/night from 1700-1900 h and 2100-2300 h, and this pattern was repeated over 2 days for each site. At the inland sites, the same protocol was followed, but the grassland transect was replaced with a second bushland transect. In addition, opportunistic collections were made at the Turkana Basin Institute (TBI) near Illeret. The localities of all individuals of reptiles and amphibians were recorded with a GPS (coordinates in latitude, longitude, datum WGS 1984, in decimal degrees), time of day was documented, and a brief Amphib. Reptile Conserv. Fig. 2. Representative examples of typical habitats within the surveyed transects in the study area: (A) cattle grazing on the grassland transect in KF; (B) bushland transect with compact soil and loose rocks and stones in KF; (C) overgrazed grassland transect at IL; (D) LO bushland transect; (E) AB bushland transect; (F) dry lagga and adjacent riverine vegetation as part of the KF river transect; (G) KA riverbed after rain; and (H) section of the IL river transect. description of the locality was given. When possible, air, substrate/water and cloacal temperatures were recorded and they are reported here. All temperature were taken by means of a K-type thermocouple (B + B Thermotechnik, effective measurement range -50 to +260 °C) connected to a digital thermometer (Center 300) (accuracy + 0.1% +0.7 °C, resolution 0.1 °C, effective measurement range -200 to +1,370 °C). Body temperatures (T,) were measured by means of cloacal temperature with the thermocouple inserted 10-15 mm into the animal’s cloaca. Taxonomy follows information from Amphibiaweb (https://amphibiaweb.org), Channing and Howell (2006), Frost (2022), Largen (2001), Uetz et al. (2022), and Spawls etal. (2018, 2019). Selected individuals were collected as voucher specimens; and when necessary, individuals were euthanized using an aqueous solution of benzocaine (20%) injected into the body cavity. After injection, the individuals were measured, a tissue sample (liver or muscle from dissected individuals, tail tips from lizards, a ventral scale clip from snakes, and toe clips from frogs) was taken and transferred to ethanol (96%) for possible future molecular analysis, and the specimen was fixed in 2% formalin. After the expedition, voucher specimens and tissue samples June 2023 | Volume 17 | Number 1/2 | e324 The Herpetofauna of Sibiloi National Park, Kenya were accessioned into the herpetology collection at NMK. For each species account, NMK numbers and/ or field numbers (for the tissue samples from voucher specimens, the specimen number is used, additional tissue samples without vouchers are listed separately), CITES status (where applicable), and any IUCN Red List status other than Least Concern are provided in the header of each species account. Results In total, we recorded 34 species, including six amphibians (Fig. 3) and 28 reptiles comprised of one freshwater turtle, one crocodile, 18 lizards, and eight snakes (Fig. 4). Many taxa in the area currently have multiple subspecies that warrant rigorous revision, so only the binomial names (genus plus species epithet) are used in this list. For each voucher specimen collected and listed here, tissue samples in 96% molecular grade ethanol are available at the NMK, and these samples can be used in future taxonomic analyses. Species Accounts Amphibia Anura Bufonidae Lugh Toad Poyntonophrynus lughensis (Loveridge, 1932) Vouchers: NMK-A1824 (field no. SK1112) Localities in the study area (henceforth simply called localities): Turkana Basin Institute (TBI) Remarks: On 28 March 2017, tens to hundreds of individuals were calling in the late morning (0900- 1100 h) together with Tomopterna wambensis in a temporary water body created by recent rainfalls in the usually dry bushland in front of the TBI near Illeret. Several pairs of Poyntonophrynus lughensis were found in amplexus. One female was collected and accessioned in the NMK collection (Fig. 3A). Similar to other members of the African pygmy toads of this genus, the natural history of P. /ughensis is almost completely unknown (Ceriaco et al. 2018). Interestingly, in a recent large-scale molecular phylogeny of African toads, P. /ughensis actually clustered with the species of the genus Mertensophryne Tihen, 1960 (Liedtke et al. 2017). Sub-desert Toad Sclerophrys xeros (Tandy, Tandy, Keith, and Duff- MacKay, 1976) Vouchers: NMK-A1817 (field no. SK1118) Localities: TBI Remarks: One male was calling on 13 April 2017 in an artificial water reservoir at TBI. This was the only individual we recorded of this widespread species from dry savanna and semi-desert. The specimen (Fig. 3B) showed the bright scarlet vermiculation on the posterior femoral integument typical for S. xeros. In addition, the call resembled typical S. xeros calls. Amphib. Reptile Conserv. Turkana Toad Sclerophrys turkanae (Tandy and Feener, 1985) IUCN Red List: Data Deficient Vouchers: NMK-A1816 (field nos. SK16 1057, SK16 1059, SK16 1060-1064) Localities: AB (R), KF (G) Remarks: Endemic to Kenya. This toad (Fig. 3C) was very abundant in the grassland transects along the shores of Lake Turkana, and found in high numbers and different sizes at Alia Bay and Koobi Fora. During the day they were hiding in moist mud cracks, and at night they were on land in slightly flooded grassy plains right at the lake edge. Males were calling in November, and the calls affirmed their identification as S. turkanae. According to the latest published amphibian guide (Spawls et al. 2019), this species is only known from Lake Turkana. The IUCN assessment mentions records from two localities in north-central Kenya: Loyangalani [Loiengalani] (south-eastern shores of Lake Turkana), and Uaso Nyiro River in the Samburu Game Reserve (IUCN SSC Amphibian Specialist Group 2016). Ptychadenidae Nile Ridged Frog Ptychadena nilotica (Seetzen, 1855) Vouchers: NMK-A1818 (field nos. SK16 1045, SK16 1046, SK16 1056, SK16 1058, SK16 1065— 1068, SK16 1070, SK16 1097, SK16 1105) Localities: IL (G), AB (R), KF (G) Remarks: This species (Fig. 3D) was found in high numbers. Individuals were found at night together with Sclerophrys turkanae in flooded grassy plains along the shore of Lake Turkana, but also on the edge of temporarily flooded waterbodies and in the riverbed transects slightly further away from the actual lake shore where S. turkanae was mostly absent. When disturbed at the water edge on land, individuals of P. nilotica escaped by jumping into the lake, but instead of diving into the water, they rather jumped across the water surface for several meters without sinking before jumping back on land. Schilluk Ridged Frog Ptychadena cf. schillukorum (Werner, 1907) Vouchers: NMK-A1825 (field no. SK16 1105) Localities: IL (R) Remarks: Ptychadena schillukorum currently has a large distribution range across sub-Saharan Africa, from Mauritania to Somalia and south to Angola and Mozambique. The taxonomy of frogs referred to as P. schillukorum is unresolved, and this taxon may actually represent a species complex (e.g., Nago et al. 2006). We assigned individual SK16 1105 (Fig. 3E) to P. cf. schillukorum based on the following features: skin of the flank granular or warty, tympanum-eye diameter ratio about 0.70, both internarial distance and nostril-tip of snout distance less than nostril-eye distance. June 2023 | Volume 17 | Number 1/2 | e324 Kirchhof et al. A previous record from the Omo River delta in Ethiopia already exists. On the night of 9 December 2016, the sole individual SK16 1105 was found sitting on soft sand in the dry riverbed of an ephemeral river (known as “lagga” or “lagha”), about 1.6 km from the lake shore. In arid lands, this species usually inhabits permanent water bodies such as swamps and springs where they can occur in sympatry with P. nilotica (e.g., in Shompole swamp, Magadi, southwestern Kenya) (PK Malonza, pers. obs.). Molecular analyses will help to resolve the taxonomic status of the P. schillukorum group. Pyxicephalidae Wamba Sand Frog Tomopterna wambensis Wasonga and Channing, 2013 Vouchers: NMK-1815 (field nos. SK16 1024, SK16 1071, SK16 1084, SK 1111) Localities: IL (G, R), KA (R), KF (R), LO (R), TBI Remarks: This medium-sized, stout, semi-fossorial frog was quite common in the study area. Its inner metatarsal tubercle is used for digging into the soil of dry riverbeds in order to reach moister areas and survive droughts, and the outer metatarsal tubercle is absent. It can be further identified by its interrupted glandular ridges below the tympanum. As soon as a few drops of rain fell — often during the night — the buried individuals of 7: wambensis (Fig. 3F) appeared on the surface. On 28 March 2017, tens to hundreds of individuals were calling in the late morning (0900-1100 h) together with Poyntonophrynus lughensis in a temporary water body created by recent rainfalls in the usually dry bushland in front of the TBI. For one individual, an internal field body temperature (T,) of 29.4 °C was recorded at an air temperature (T,) of 30 °C and substrate temperature (T,,) of 21 °C. Reptilia Testudines Trionychidae Nile Soft-shelled Turtle Trionyx triunguis (Forskal, 1775) IUCN Red List: Vulnerable. CITES App. I Vouchers: None Localities: KF (B) Remarks: The carapace of one dead individual of this soft-shell turtle was found about 900 m from the shore of Lake Turkana at Koobi Fora. These turtles are still eaten by the local people around Lake Turkana (V. Wasonga, pers. obs.; Spawls et al. 2018), so we suspect the locality does not reflect the individual’s actual habitat. The Nile Soft-shelled Turtle inhabits permanent lakes, dams, and rivers, and is known to enter the sea, but it does not live in any other Kenyan lake in the Great Rift Valley other than Lake Turkana, because none of the other rivers were ever connected to the Nile system (Spawls et al. 2018). Crocodylia Crocodylidae Nile Crocodile Crocodylus niloticus Laurenti, 1768 CITES App. I Vouchers: None Localities: AB (L), IL (L, B), KF (L, G) Remarks: The Lake Turkana population of Nile Crocodiles used to be the largest in the world (IUCN World Heritage Outlook 2020). Recent data suggests that the crocodiles are heavily impacted by local fishermen who destroy nesting sites, and the remaining population is only a fraction of what it used to be (UCN World Heritage Outlook 2020). We found Crocodylus niloticus in Lake Turkana and adjacent water bodies along the shoreline. Very young juveniles (Fig. 4A) were found in March/April. Fig. 3. Amphibian species recorded during the surveys: (A) Poyntonophrynus lughensis, (B) Sclerophrys xeros; (C) Sclerophrys turkanae, (D) Ptychadena nilotica, (E) Ptychadena cf. schillukorum;, and (F) Tomopterna wambensis. Amphib. Reptile Conserv. June 2023 | Volume 17 | Number 1/2 | e324 The Herpetofauna of Sibiloi National Park, Kenya Squamata Agamidae Red-headed Rock Agama Agama lionotus Boulenger, 1896 Vouchers: NMK-398L/1 (SK16 1038b), 399L/1-—2 (field nos. SK16 1114-1115) Distribution: AB (R), petrified forest Remarks: The distribution of this taxon is still uncertain in many parts of its range because of possible confusion with Agama agama (but not in Kenya), A. finchi, and A. dodomae (Spawls et al. 2018). This agama was only found in Alia Bay (three individuals including one juvenile NMK-398L/1 (SK16 1038b)) and in the petrified forest (NMK SK1115). In the absence of rocks, this species was entirely arboreal, living on Vachellia sp. trees in the riparian woodlands along the dry riverbeds. Only in the rock-dominated petrified forest site (about 5 km from Karsa Gate, the southern entry point of SNP) was this species rupicolous. We found Agama lionotus individuals (Fig. 4B) only between 1800 h and 2100 h. The average T, was 33.8 + 1.7 °C (32.6-35 °C; N = 2) with T., ranging from 30-34 °C and T, ranging from 32-35 °C. Riippell’s Agama Agama rueppelli Vaillant, 1882 Vouchers: NMK-375L/1-4 (field nos. SK16 1042, SK16 1074, SK16 1075, SK16 1094) Distribution: AB (B), KA (B, R), KF (R), LO (B) Remarks: This species occurs in dry savannas and semi- deserts in lower elevations (Spawls et al. 2018). We recorded 25 individuals of this strictly terrestrial agama (Fig. 4C) in bushland at all sites except IL. Activity was high during all four study months, and individuals were observed to be active from 0900 h to 2200 h. The average T, was 31.9 + 3.0 °C (28.4-36.5 °C; N = 5) with T., ranging from 24-39 °C and T, ranging from 27—36 °C. b Eublepharidae Somali-Maasai Clawed Gecko Holodactylus africanus Boettger, 1893 Vouchers: NMK-390L/1-3 (field nos. SK16 1080, SK16 1081, SK16 1083) Distribution: KA (R), IL (R), TBI Remarks: Very little is known about this nocturnal terrestrial eyelid gecko and only occasional records exist. It is assumed to occur throughout dry savannas and semi-deserts from southeastern Ethiopia, northern Somalia through Kenya and into northeastern Tanzania at low elevations of 200—1,000 m (Spawls et al. 2018). We found seven individuals in November, December, and April in dry sandy river beds at KA and at TBI near the water reservoir. The species’ occurrence seems to be associated with high groundwater and/or dense vegetation as all individuals were found at night in the riverbed at the edge of the vegetated slope. Three different color morphs were found, including yellowish, slightly pink, and darker brown (Fig. 4D). These are the first Kenyan records from the eastern side of Lake Turkana. Amphib. Reptile Conserv. Gekkonidae East African House Gecko Hemidactylus angulatus Hallowell, 1852 Vouchers: NMK-380L/1-3 (field nos. SK16 1110, SK 1116, SK16 1055) Localities: KF (B), TBI, petrified forest Remarks: This is a fairly large gecko that is widespread in Kenya, occurring south to Tanzania, north to Sudan and west to Senegal, and occupying a variety of dry and mesic habitats (Spawls et al. 2018). It can be distinguished from H. ruspolii and other Hemidactylus by small granules covering the upper surface of the snout rather than large and keeled granules, heterogeneous dorsal scales with small granular scales interspersed with rows of large, keeled, and mainly oval tubercles, and males with a series of 20-46 preano-femoral pores interrupted mid-ventrally (Fig. 4E). Three individuals were found over the course of our study, and all of them were associated with rocks or anthropogenic structures/ houses. This taxon might actually represent a species complex (the type locality is in Gabon), and extensive molecular analyses are necessary to resolve its status. One gravid female carrying eggs was collected on 7 April 2017. Barbieri’s Turkana Gecko Hemidactylus barbierii Sindaco, Razzetti, and Ziliani, 2007 Vouchers: NMK-378L/1—4 (field nos. SK16 1078, SK16 1085—1087) Distribution: KA (R) Remarks: Four individuals were found at KA (about 15 km inland of Lake Turkana) at about 514 m elevation. These represent the first records from the eastern side of Lake Turkana (Sindaco et al. 2007). This species can be distinguished from the syntopic Hemidactylus ruspolii Boulenger, 1896 by the presence of precloacal pores in males (instead of femoral pores), and a dorsal pattern consisting of four transverse dark grey (or at least bordered with dark grey margins) bands (one nuchal and three between anterior and posterior limbs), and a dark rather narrow crescent shaped band bordering the posterior skull margins and extending through the eyes to the nostril (more pronounced in the yellow and black juveniles; Fig 4F). It is different from Hemidactylus bavazzanoi Lanza, 1978 in the mental scale arrangement as well as the dorsal pattern, which in H. bavazzanoi comprises only three dark transverse bands and a broader crescent shaped head band. Little is known about the ecology of H. barbierii. It is usually considered to be terrestrial, hiding under ground coverings or in holes (Sindaco et al. 2007; Spawls et al. 2018). Based on our observations, we consider it to be arboreal rather than terrestrial. This species inhabits the riparian woodlands along dry laggas where individuals were found actively foraging at night on the tree stems and at the bases of trees. All individuals escaped by fleeing up the trees rather than running away on the ground. One recently hatched individual (Fig. 4F) was found during the day hiding under the bark of a Vachellia sp. tree at about 1.8 m above the ground on 29 November 2019. June 2023 | Volume 17 | Number 1/2 | e324 Kirchhof et al. > aagsek ai 2 sem : Pee. Fig. 4. Reptile species recorded during the surveys: (A) Crocodylus niloticus, (B) Agama lionotus,; (C) Agama rueppelli, (D) Holodactylus africanus; (EK) Hemidactylus angulatus;, (F) Hemidactylus barbierii; (G) Hemidactylus lanzai, (H) Hemidactylus ruspolii; (1) Homopholis fasciata;, (J) Lygodactylus somalicus, (1K) Stenodactylus sthenodactylus, (L) Heliobolus spekii; (M) Latastia longicaudata; (N) Philochortus rudolfensis, (O) Chalcides bottegi; (P) Mochlus sundevallii, (Q) Trachylepis striata; (R) Varanus albigularis; (S) Eryx colubrinus, (T) Platyceps brevis, (U) Psammophis cf. tanganicus;, (V) Psammophis punctulatus, (W) Rhamphiophis rostratus, (X) Naja pallida; (Y) Bitis arietans, and (Z) Echis pyramidum. Amphib. Reptile Conserv. vA June 2023 | Volume 17 | Number 1/2 | e324 The Herpetofauna of Sibiloi National Park, Kenya So far, this species 1s only known from the Lake Turkana region. The type series was collected south of KF, 10 km inland from AB (Sindaco et al. 2007). Lanza’s Gecko p Hemidactylus lanzai Smid, Mazuch, Novakova, Modry, Malonza, Elmi, Carranza, and Moravec, 2020 et al. 2020 Vouchers: NMK-391L (field no. SK16 1021); NMK- 383L/1-2 (field nos. SK16 1029, SK16 1033); NMK-395L (field no. SK16 1039); NMK-392L (field no. SK16 1053); NMK-393L (field no. SK16 1037) Additional tissue samples: SK105 2016 Localities: AB (R), IL (R, G, B), KA (R, B), KF (R, G), LO (R) Remarks: Recently elevated to a full species from the synonymy of its sister species Hemidactylus isolepis Boulenger, 1895, this small, terrestrial gecko is a typical inhabitant of the dry, semi-desert areas in northern Kenya (Spawls et al. 2018). This species (Fig. 4G) was one of the most common reptiles in SNP, with 96 individuals recorded. The highest abundance was along the shores of Lake Turkana, where individuals were hiding in the root system under dried reeds, but it also was found tn dry laggas and bushland under dead logs, trees, and rocks. It was exclusively active at night, foraging on sand, gravel, and compact soils. The average T, was 34.0 + 1.5 °C (30.5-35.8 °C; N = 10) with T. , ranging from 24-47 °C and T, ranging from 28-38 °C. Prince Ruspoli’s Gecko Hemidactylus ruspolii Boulenger, 1896 Vouchers: NMK-377L/1—11 (field nos. SK16 1026, SK16 1028, SK16 1032, SK16 1040, SK16 1044, SK16 1054, SK16 1073, SK16 1089) Localities: AB (R), IL (B, R), KA, (B, R), KF (B, R) Remarks: This medium-sized gecko (Fig. 4H) is less widespread in East Africa and inhabits drier and lower areas in Kenya, Somalia, and Ethiopia compared to its relative H. angulatus. This species was quite abundant in the riverbeds of SNP were we often found it at night under bark, foraging along tree stems, and at the foot of trees. A few individuals were also found in bushland on Salvadora persica (“toothbrush tree”). Average T, was 32.3 + 1.0 °C (30.5-34.1 °C; N= 12) with T. , ranging from 25—33 °C and T, ranging from 29-34 °C. Banded Velvet Gecko Homopholis fasciata (Boulenger, 1890) Vouchers: NMK-386L/1-—2 (field nos. SK16 1095, SK16 1103) Localities: AB (R), IL (R), KA (B), LO (R) Remarks: The strictly arboreal Banded Velvet Gecko (Fig. AT) is a savanna species that typically occupies holes and crevices in large trees from sea level to at least 1,300 m (Spawls et al. 2018). In SNP, we found six individuals in December, March, and April. Individuals were found under bark and in holes of Vachellia sp., as well as on the much smaller and bush-like Salvadora persica. A gravid female was collected on 30 March 2017. The average T, was 35 + 0 °C (N =2) with T. , ranging from 33—34 °C and T, ranging from 34—35 °C. These are the first Kenyan records from the eastern side of Lake Turkana. Amphib. Reptile Conserv. Somali Dwarf Gecko Lygodactylus somalicus Loveridge, 1935 Vouchers: NMK-387L/1-3 (field nos. SK16 1072, SK16 1101, SK16 1104) Additional tissue samples: SK278 2016 Localities: IL (R, B), KF (R) Remarks: This arboreal species is a typical inhabitant of dry savannas and semi-deserts. During our surveys, 39 individuals of this small diurnal gecko (Fig. 4J) were recorded in IL (N = 35) and KF (N = 4), in both bushland and in dry riverbeds on small trees and shrubs (the majority on Salvadora persica, but also on Vachellia sp.). Elegant Gecko Stenodactylus sthenodactylus (Lichtenstein, 1823) Vouchers: NMK-396L/1-3 (field nos. SK16 1022, SK16 1030, SK16 1031) Additional tissue samples: SKO84 2016, SK120 2016, SK117 2016 Localities: AB (R, B), IL (R, B, G), KA (B), KF (R, B, G), LO (B) Remarks: This widespread nocturnal, terrestrial gecko (Fig. 4K) is known from semi-deserts and deserts across its range, but in Kenya it has only been recorded in the Lake Turkana vicinity. It was very common in the study area, with 75 individuals recorded. All individuals were found on the ground (sand, gravel, often under shrubs) in the evening and at night (1900 h to 2300 h). The average T, was 32.4 + 2.2 °C (25.8-35 °C; N = 20) with T,,, ranging from 23-33 °C and T, ranging from 27-35 °C. Lacertidae Speke’s Sand Lizard Heliobolus spekii (Ginther, 1872) Vouchers: NMK-376L/1-3 (field nos. SK16 1076, SK16 1082, SK16 1088) Additional tissue samples: SK351 2017, SK468 2017, SK469 2017, SK478 2017, SK479 2017 Localities: KA (B, R), LO (B), TBI Remarks: Heliobolus spekii (Fig. 4L) is a widespread generalist species inhabiting coastal thicket and woodland, moist and dry savanna, and semi-desert at elevations from sea level to 1,500 m (Spawls et al. 2018). We found 18 individuals throughout the study period. This species was absent from the sites near the lake shore and from sparsely vegetated areas. It was only found further inland at the Karare site, Lomosia, and at TBI in more densely vegetated, grassy bushland and on vegetated riverbanks. One pair was observed mating on 17 April 2017. The average T, was high at 37.5 + 1.4 °C (34.5-39 °C; N = 8) at substrate temperatures (T.,) ranging from 28-49 °C and T, ranging from 29-34 °C. Long-tailed Sand Lizard Latastia longicaudata Reuss, 1834 Vouchers: NMK-388L (field no. SK16 1027); NMK-385L (field no. SK1034); NMK-384L (field no. SK1050) Additional tissue samples: SK110.1 2016, SK142 2016, SK158 2016, SK195 2016, SK233 2016, SK295 2017, SK303 2017, SK305 2017, SK306 2017, SK352 2017, SK481 2017 June 2023 | Volume 17 | Number 1/2 | e324 Kirchhof et al. Localities: AB (R, B, G), IL (R, B, G), KA (R, B), KF (R, B), LO (R, B) Remarks: We recorded 33 individuals of this common diurnal lacertid (Fig. 4M) throughout the study period on all transects between 0800 h and 1930 h. All lizards were darting between bushes in search of prey and places for hiding. Average T, was high at 37.8 + 1.2 °C (36.2— 39.5 °C; N = 8) at T.,, ranging from 31-65 °C and T, ranging from 33-39 °C. The genus Latastia consists of multiple similar-looking species and subspecies, and in some cases, only the suspected distribution range allows for identification. Rigorous genetic analysis 1s necessary to update the taxonomic status, the distinguishing morphological character traits, and the distribution ranges of existing specimens. Turkana Shield-backed Ground Lizard Philochortus rudolfensis Parker, 1932 Vouchers: NMK-382L/1-2 (field nos. SK16 1048, SK89 2016) Localities: AB (B) Remarks: Only a limited number of records exist for this species, and its exact range is therefore unknown -— it 1s currently known from southern Ethiopia and northern and central Kenya, at elevations below 800 m (Spawls et al. 2018). During our survey, we found two individuals (Fig. 4N) in a very dry and overgrazed area of the AB bushland on compacted but not rocky substrate in the late mornings of 15 and 16 November 2016. On the same transect, Agama rueppelli, Latastia longicaudata, Rhamphiophis rostrastus, Stenodactylus sthenodactylus, and Trachylepis striata were also recorded. Scincidae Ocellated Skink Chalcides bottegi Boulenger, 1898 Vouchers: NMK-389L (field no. SK16 1092) Localities: KA (B), KF (G) Remarks: This skink (Fig. 40) is known from dry savanna and semi-desert in northern Kenya. We found only two individuals of this semi-fossorial skink in KA and KF in grassland and bushland, respectively. Both individuals were active during the day, one was found under dead reeds along the lake shore, and the other was digging into the loose sand under a Commiphora africana shrub. The body temperature of the individual under the reed was 34.9 °C at a substrate temperature of 34 °C (T, 37.5 °C). These are the first Kenyan records from the eastern side of Lake Turkana. Sundevall’s Writhing Skink Mochlus sundevallii (Smith, 1849) Vouchers: NMK-397L/1-—4 (field nos. SK16 1035, SK16 1077, SK16 1090, SK16 1106) Additional tissue samples: SK281 2016 Localities: IL (B, R), KA (R), KF (R, B, G), LO (B) Remarks: A nocturnal, fossorial species that occupies a variety of habitats comprising coastal savanna and woodland, dry and moist savanna, semi-desert, and medium to high-elevation woodland up to 2,000 m (Spawls et al. 2018). Seventeen individuals of Mochlus Amphib. Reptile Conserv. sundevallii (Fig. 4P) were found, all after sunset, digging in soft sand under shrubs. Average T, was 33.9 + 1.0 °C (32.7-34.9 °C; N = 4) at T., ranging from 29-30 °C and T, ranged from 32-35 °C. Striped Skink Trachylepis striata (Peters, 1844) Vouchers: NMK-379L/1-—5 (field nos. SK16 1036, SK16 1041, SK16 1052, SK16 1069, SK16 1107) Localities: AB (B, L), IL (B), KA (B), KF (G) Remarks: A recent study found Trachylepis striata (Fig. 4Q) to be paraphyletic, with Ethiopian individuals being the sister group to Tanzanian individuals and 7? mlanjensis (Loveridge, 1953) from Malawi (Weinell et al. 2019). Kenyan specimens were not included in that study. Extensive molecular analyses of individuals covering the distribution ranges of these taxa will be necessary to resolve their respective status. For now, Trachylepis striata is considered a generalist skink, living in forest clearings, coastal thicket, moist and dry savanna, semi-desert, and urban areas (Spawils et al. 2018). The 14 individuals of a diurnal skink found during this study were assigned to 7’ striata sensu lato. All individuals were discovered on the ground, although this species has been referred to as arboreal (Spawls et al. 2018). The majority of individuals were found under clumps of dead reeds along the lake shore. Average T, was 34.7 + 0.4 °C (34.4-35 °C; N = 2) at T., ranging from 34-40 °C and T, was 34 °C, Varanidae Savannah Monitor Varanus albigularis Daudin, 1802 CITES App. II Vouchers: NMK-381L (field no. SK16 1108) Localities: IL (R), AB (R) Remarks: This monitor lizard 1s quite widespread across Africa and occupies different habitats from dry and moist savanna, coastal thicket and woodland, and semi-desert, from sea level to 1,500 m elevation (Spawls et al. 2018). It likely occurs across most of Kenya, but museum specimens are scarce. The records of the rock or White-Throated Monitor represent the first records for the eastern shore of Lake Turkana, although there was hardly any doubt the species occurred there. Three individuals were found in December and April, comprising one adult and two juveniles. The two juveniles (Fig. 4R) were found at night sleeping on tree branches, one was exposed just lying atop the branch at 2 m height, and one was hidden under bark close to the ground. The adult individual was found in the morning under the bark of a tree stem. Nile Monitor Varanus niloticus (Linnaeus, 1766) CITES App. I Vouchers: None Localities: IL (G) Remarks: The most widespread African lizard usually lives near fresh water sources from sea level to around 1,600 m elevation, and rarely higher. We observed one individual Nile Monitor walking along the shore of Lake June 2023 | Volume 17 | Number 1/2 | e324 The Herpetofauna of Sibiloi National Park, Kenya Turkana in the early afternoon in December 2016. This species might be under threat in some parts of Africa, as it is exploited for its skin, particularly in West Africa, but the proliferation of dams has provided extra habitat for this species (Spawls et al. 2018). It is probably quite common in SNP. Serpentes Boidae Kenya Sand Boa Eryx colubrinus (Linnaeus, 1758) CITES App. II Vouchers: NMK-372S (field no. SK16 1023) Additional tissue samples: SK353 2017 Localities: AB (B), IL (B, R), KF (B), LO (B), TBI Remarks: A typical inhabitant of arid and semi-arid areas in north-eastern Africa (Spawls et al. 2018). The natural history of the Sand Boa (Fig. 4S) is poorly known due to its secretive fossorial lifestyle. We recorded seven individuals. Six of them were active at night or at least after sunset, and the other one recorded during daytime was found at TBI at noon after rain. It was a large female that was lying under a tree in the moist soil with its head buried underground and its entire body exposed; however, it was largely protected from direct sun by the shadow of the tree. Individuals were recorded in November, March, and April mainly in bushland, and only one individual was found in a dry riverbed. Colubridae Smith’s Racer Platyceps brevis (Boulenger, 1895) Vouchers: NMK-373S (field no. SK16 1049) Localities: AB (house wall) Remarks: This racer occupies dry savanna and semi-desert at elevations spanning 100—1,300 m (Spawls et al. 2018). In the early afternoon (1400 h) of 16 November 2016, we collected a specimen of this species (Fig. 4T) coming out of a crevice in the wall of one of the buildings in the Alia Bay grasslands. Only limited information is known about this species and records are still sparse, leading to a fragmented currently-known distribution range (Spawls et al. 2018). Psammophiidae Tanganyika Sand Snake Psammophis cf. tanganicus Loveridge, 1940 Vouchers: NMK-370S (field no. SK16 1093) Records: KA (B) Remarks: A slim grey sand snake very similar to Psammophis biseriatus, of which it was originally described as a subspecies. Its taxonomic status will remain unresolved without thorough genetic and morphological analyses covering their entire ranges. According to Loveridge (1940) the only character separating P. biseriatus biseriatus from P. b. tanganicus [sic] is the number of labial scales entering the orbit (two in P. b. biseriatus vs. three in P. b. tanganicus [sic]). Specimen NMK-370S has nine labials and the 4", 5", and 6" are Amphib. Reptile Conserv. in contact with the orbit. Labials are not plain white but rather largely blotched in light brown. This poorly known snake occurs from sea level to about 1,300 m in dry savanna and semi-desert, but the known Kenyan records are few and very scattered (Spawils et al. 2018). We found only one individual (Fig. 4U) in the bushland at Karare in the evening of 2 December 2016 right at sunset. It was actively moving on the ground, but immediately climbed into a shrub when disturbed. If the specimen proves to be P. tanganicus this will be the first record from the Lake Turkana area. Genetic analyses are necessary to resolve the taxonomic status of P. biseriatus and P. tanganicus. Speckled Sand Snake Psammophis punctulatus Duméril, Dumeéril, 1854 Vouchers: NMK-S4604 (field no. SK1113) Records: AB (R), KF (museum headquarters) Remarks: This common diurnal snake is widely distributed. We found one adult and one juvenile individual in Koobi Fora and Alia Bay in the vegetation along the riverbed and in the Koobi Fora camp, both in April. The large adult individual (Fig. 4V) was feeding on a weaver bird when found (Kirchhof et al. 2018). The currently accepted subspecies P. p. punctulatus and P. p. trivirgatus (to which the SNP specimens were assigned) seem to be parapatric and morphologically distinguishable, and they might both prove to be full species in the future. Bibron, and Rufous Beaked Snake Rhamphiophis rostratus Peters, 1854 Vouchers: NMK-368S (field no. SK16 1051) Localities: AB (B) Remarks: This diurnal snake inhabits semi-desert, dry and moist savanna, coastal thicket, and woodland up to 1,500 m (Spawls et al. 2018). We recorded this large terrestrial snake only once (Fig. 4W) in bushland at Alia Bay in November 2016 at night. Elapidae Red Spitting Cobra Naja pallida Boulenger, 1896 Vouchers: NMK-367S (field no. SK16 1047) Localities: KA (R), KF (G, R) Remarks: This spitting cobra is another typical occupant of semi-deserts and dry savannas (Spawils et al. 2018). In November 2016, we found three individuals of the gray- phased Naja pallida (Fig. 4X). All were active after sunset at Koobi Fora and Karare. One was found in grassland near the lake shore, and the other two were foraging in dry riverbeds. Viperidae Puff Adder Bitis arietans (Merrem, 1820) Vouchers: NMK-369S (field no. SK16 1102) Localities: IL (R) Remarks: We recorded one individual (Fig. 4Y) of this large and widely distributed viper at night in December 2016 among the Vachellia sp. trees in a dry riverbed. June 2023 | Volume 17 | Number 1/2 | e324 Kirchhof et al. ‘TROL, ‘JOL, ‘pueyysng ‘g {poqioAl “Y Spuvjssess “H “eIsowo’T ‘—O'T ‘dW yI] “TI ‘resIe yy ‘({B1OL, JOT, ‘pueyysng “g “paqJoAls “Y Spuvylsseis DH ‘VY ‘e10y Iqooy “Fy SAeg PITY “GV :SuUOneIADIQqY ‘S}OOSUBI] DY} JO 9UO JO SoS POADAINS “D) syoosuvl] pue (VISOWO'T ‘O’'T ‘dWIOyI] “TI ‘esey “vy “eso, Iqooy “py ‘Aeg eITV bel dy} JO QUO JOYS ye puNOJ ATSAISN[OXS 919M Jey) Sorsods uviqrydue Jo sSIOqUINN °g “SITY “qV) sous ADAINS OY} SSOIOB SoIdads ULIqIydWe papsOSdI XIS dy} JO SDUDIINIIO *L “SI my aus Apnig ays Apnis 5 yasues, dwey lal O1 al) WA dy av WLOL r= 3 : ; ‘ r 9 0 | 0 [ 0 fn) I i) i 0 ) | 0 i ) : 3 q 2 g q 9 q q 2) q me, a o1 WH av : N T T T T T T T @ 4 4OL Y 4OL uy | yoL = LZ = ee Zz z Zz S T T = uo 9) 3 —_ 11 4 3 z QI io” € oO N g € € € € 3 a iF }OL yO, }OL 5 o oO % i = 8 > me 32 +28 ¥ i z lat r gs 9 9 € 30] Ad, ‘TROL ‘OL -pueyysnq : (4 ¢ (4 ¢ (4 ¢ (4 5 ‘q ‘paqiaall “Y ‘puvyssess “H ‘anjnsuy ulseg eueYIN], ‘Tq I “eIsowoy ‘OT ‘a1oway]] “TI ([e10], JOL, :puerysng “g ‘poqioals “Y :pueysses “H) Joosuey Aq pue (o)NIVsSU] ‘oIVIey “WY ‘B10 Iqooy “Py ‘Avg PITY “GV ‘SUOTIEIADIQGY ‘s}oosUe dU} JO UO IO SaqIS ieee Ye TEL Bou TO 1 a te TT eal Wake ed OS ae ‘Keg poAdAINs dy} JO DUO JOYITA ye PUNO} AJAAISNOXS d19M Je} Sar1dads ayd91 Jo SIOQUINN *9 “SIH BITV “@V) Sous AoArns oy} ssosoe sarseds a]dos pops0dE1 BZ SY} JO S9USIINIIO *¢ “SIA ays ApN|S ays Apis ial Wy WILOL pesuedl dwey ! a Zz : = 3 © ion = ® gS 3 OL Q, n ie) in = 2. 10) 10] JO] oH ro) o 10] ) 2 = 2 ® a re ic E Ls yo, (OE xt The Herpetofauna of Sibiloi National Park, Kenya Table 1. List of the amphibian and reptile taxa recorded at Sibiloi National Park, and additional sites along the eastern shore of Lake Turkana south to Mount Kulal, during the present study and from Ziliani et al. (2016). The list does not include Psammophis semivariegatus, which is listed by Ziliani et al. (2006), as this taxon does not exist. We assume that this name refers to Philothamnus semivariegatus (Smith, 1840) which was listed as Philothamus semivariegatus. We here use the species names as used in Ziliani et al. (2016). Some of these names have undergone taxonomic changes by now, for others we could not find out which taxon was referred to (indicated with (?)), and sometimes the taxon name was misspelled (indicated with [sic]). This study Ziliani et al. (2006)* AMPHIBIA Bufonidae Poyntonophrynus lughensis (Loveridge, 1932) x Bufo lughensis Sclerophrys xeros (Tandy, Tandy, Keith, and Duff-MacKay, 1976) x - Sclerophrys turkanae (Tandy and Feener, 1985) x Bufo turkanae Sclerophrys cf. gutturalis (Power, 1927) = Bufo cf. gutturalis (Mt. Kulal) Ptychadenidae Ptychadena nilotica (Seetzen, 1855) x Ptychadena mascareniensis (?) Ptychadena anchietae (Bocage, 1868) — Ptychadena anchietae Ptychadena cf. schillukorum (Werner, 1907) x — Ptychadena cf. taenioscelis Laurent, 1954 — Ptychadena cf. taenioscelis Pyxicephalidae Tomopterna wambensis Wasonga and Channing, 2013 x Tomopterna cryptotis (?) REPTILIA Pelomedusidae Pelomedusa neumanni Petzold, Vargas-Ramirez, Kehlmaier, Pelomedusa subrufa Vamberger, Branch, Du Preez, Hofmeyr, Meyer, Schleicher, Siroky, — and Fritz, 2014 Pelusios broadleyi Bour, 1986 = Pelusios broadleyi Testudinidae Malacochersus tornieri (Siebenrock, 1903) = Malacochersus tornieri (Mt. Kulal) Trionychidae Trionyx triunguis (Forskal, 1775) 4 Trionyx triunguis Crocodylidae Crocodylus niloticus Laurenti, 1768 x Crocodylus niloticus Agamidae Agama lionotus Boulenger, 1896 x Agama agama lionotus Agama rueppelli Vaillant, 1882 x Agama rueppelli Chamaeleonidae Trioceros bitaeniatus (Fischer, 1884) = Chamaeleo bitaeniatus (Mt. Kulal) Trioceros narraioca (Necas, Modry, and Slapeta, 2003) = Chamaeleo narrayioca [sic] (Mt. Kulal) Eublepharidae Holodactylus africanus Boettger, 1893 x - Gekkonidae Hemidactylus angulatus Hallowell, 1852 xe Hemidactylus brooki (?) Hemidactylus barbierii Sindaco, Razzetti, and Ziliani, 2007 x Hemidactylus n. sp. Hemidactylus lanzai Smid, Mazuch, Novakova, Modry, Malonza, Elmi, Hemidactylus isolepis Carranza, and Moravec, 2020 * Hemidactylus macropholis Boulenger, 1896 = Hemidactylus macropholis Hemidactylus platycephalus Peters, 1854 = Hemidactylus platycephalus Amphib. Reptile Conserv. 12 June 2023 | Volume 17 | Number 1/2 | e324 Kirchhof et al. Table 1 Continued. List of the amphibian and reptile taxa recorded at Sibiloi National Park, and additional sites along the eastern shore of Lake Turkana south to Mount Kulal, during the present study and from Ziliani et al. (2016). The list does not include Psammophis semivariegatus, which is listed by Ziliani et al. (2006), as this taxon does not exist. We assume that this name refers to Philothamnus semivariegatus (Smith, 1840) which was listed as Philothamus semivariegatus. We here use the species names as used in Ziliani et al. (2016). Some of these names have undergone taxonomic changes by now, for others we could not find out which taxon was referred to (indicated with (?)), and sometimes the taxon name was misspelled (indicated with [sic]). This study Ziliani et al. (2006)* Hemidactylus ruspolii Boulenger, 1896 x Hemidactylus ruspolii Homopholis fasciata (Boulenger, 1890) xX - Lygodactylus keniensis Parker, 1936 _ Lygodactylus kenyensis [sic] Lygodactylus somalicus Loveridge, 1935 x Lygodactylus cf. somalicus Stenodactylus sthenodactylus (Lichtenstein, 1823) x Stenodactylus sthenodactylus Gerrhosauridae Gerrhosaurus flavigularis Wiegmann, 1828 = Gerrhosaurus flavigularis (Mt. Kulal) Lacertidae Heliobolus spekii (Gunther, 1872) xX Heliobolus spekii Latastia longicaudata Reuss, 1834 x Latastia longicaudata Philochortus rudolfensis Parker, 1932 x - Pseuderemias smithi (Boulenger, 1895) — Pseuderemias smithi Scincidae Panaspis sp. — Afroblepharus sp. Chalcides bottegi Boulenger, 1898 x Chalcides ocellatus bottegi Mochlus sundevallii (Smith, 1849) J Lygosoma sundevalli [sic] and Lygosoma afrum (Mt. Kulal) Trachylepis quinquetaeniata (Lichtenstein, 1823) eS Mabuya quinquetaeniata Trachylepis striata (Peters, 1844) x Mabuya striata Trachylepis varia (Peters, 1867) = Mabuya varia Varanidae Varanus albigularis Daudin, 1802. x - Varanus niloticus (Linnaeus, 1766) x — Atractaspididae Aparallactus lunulatus (Peters, 1854) = Apparalactus lunulatus [sic] (Mt. Kulal) Boidae Eryx colubrinus (Linnaeus, 1758) x Eryx colubrinus Colubridae Crotaphopeltis hotamboeia (Laurenti, 1768) 4 Crotaphopeltis hotamboeia (Mt. Kulal) Dasypeltis scabra (Linnaeus, 1758) = Dasypeltis scabra (Mt. Kulal) Dispholidus typus (Smith, 1829) = Dispholidus typus (Mt. Kulal) Philothamnus semivariegatus (Smith, 1840) 2 Philothamus semivariegatus [sic] (Mt. Kulal) Platyceps brevis (Boulenger, 1895) x Platyceps brevis smithi Platyceps florulentus (Geoffroy Saint-Hilaire, 1827) = Platyceps florulentus florulentus Telescopus obtusus (Reuss, 1834) — Telescopus dhara Elapidae Naja haje (Linnaeus, 1758) = Naja haie [sic] (Mt. Kulal) Naja pallida Boulenger, 1896 x Naja pallida Amphib. Reptile Conserv. 13 June 2023 | Volume 17 | Number 1/2 | e324 The Herpetofauna of Sibiloi National Park, Kenya Table 1 Continued. List of the amphibian and reptile taxa recorded at Sibiloi National Park, and additional sites along the eastern shore of Lake Turkana south to Mount Kulal, during the present study and from Ziliani et al. (2016). The list does not include Psammophis semivariegatus, which is listed by Ziliani et al. (2006), as this taxon does not exist. We assume that this name refers to Philothamnus semivariegatus (Smith, 1840) which was listed as Philothamus semivariegatus. We here use the species names as used in Ziliani et al. (2016). Some of these names have undergone taxonomic changes by now, for others we could not find out which taxon was referred to (indicated with (?)), and sometimes the taxon name was misspelled (indicated with [sic]). Lamprophiidae Boaedon fuliginosus (Boie, 1827) Lycophidion sp. Leptotyphlopidae Myriopholis macrorhyncha (Jan, 1860) Psammophiidae Psammophis biseriatus (Peters, 1881) Psammophis punctulatus Duméril, Bibron, and Dumeéril, 1854 Psammophis cf. tanganicus Loveridge, 1940 Rhamphiophis rostratus Peters, 1854 Rhamphiophis rubropunctatus (Fischer, 1884) Viperidae Bitis arietans (Merrem, 1820) Echis pyramidum (Geoffroy Saint-Hilaire, 1827) North-east African Carpet Viper Echis pyramidum (Geoffroy Saint-Hilaire, 1827) Vouchers: NMK-374S (field no. SK16 1025); NMK-371S (field no. SK16 1149) Additional tissue samples: SKO51 2016 Localities: AB (B, R), IL (B, R), KA (B), KF (B, G, R), LO (R) Remarks: The most common snake in SNP during our expeditions was Echis pyramidum, with 38 individuals recorded. This species (Fig. 4Z) occurred at all our study sites and in all transect types, but with a predilection for bushland (26 records). Activity was restricted to the night time, and records during daytime were exclusively of resting individuals dug out under dead logs. Camp and Transect Comparisons Of the 28 reptile species, 11 were recorded in the grasslands along the lake shore (of those, P brevis was not recorded in natural habitat but on the wall of a building), 19 in the dry riverbeds, and 21 in bushland. The species diversity of the different sites were very similar (16-17 species per site) when the sites with similar collecting effort were compared, 1.e., Lomosia with 10 species was only surveyed in March-April 2017, and species at TBI were recorded opportunistically (Fig. 5). The species found only at one site comprised four at Alia Bay (Agama lionotus, Philochortus rudolfensis, Platyceps brevis, Rhamphiophis rostratus), One species at Koobi Fora (7rionyx triunguis), two species at Karare (Hemidactylus barbierii, Psammophis cf. tanganicus), and two at Ilkemere (Bitis arietans, Varanus niloticus) (Fig. 6). Philochortus rudolfensis, P. Amphib. Reptile Conserv. This study Ziliani et al. (2006)* = Lamprophis fuliginosus (Mt. Kulal) — Lycophidion sp. (Mt. Kulal) = Leptotyphlops machrorhynchus _ Psammophis biseriatus (Mt. Kulal) x Psammophis cf. punctulatus x J 3 / Rhamphiophis rubropunctatus 4 Bitis arietans x Echis pyramidum cf. tanganicus, and R. rostratus (and the Trionyx triunguis carapace) were found only on bushland transects; A. lionotus, B. arietans, Hemidactylus barbierii, and Varanus albigularis occurred exclusively in dry riverbeds; and P. brevis and V. niloticus were only found in the grassland transects at IIkemere and Alia Bay, respectively (Fig. 6). Most of the six species of amphibians (four species; Ptychadena nilotica, Ptychadena cf. schillukorum, Sclerophrys turkanae, and Tomopterna wambensis; Fig. 7) were found in the dry riverbeds, especially after rains. Three amphibian species were recorded in grasslands. Of those, P. nilotica and S. turkanae were found in the highly alkaline (pH = 9.2) and saline (TDS = 2,500 ppm; Yuretich and Cerling 1983) water of Lake Turkana. The fossorial 7. wambensis either appeared from out of its underground hiding place away from the waters after rain, or it was calling at sites away from the lake at the edges of confined water bodies, which were most probably fed mainly by the high levels of groundwater and rain water. Of the three species recorded at TBI, 7) wambensis and Poyntonophrynus lughensis were mating in a freshly flooded temporary waterbody after heavy rains in bushland, while Sclerophrys xeros was sitting in the artificial water reservoir of the station. Only a single individual of P. cf. schillukorum was found in a dry riverbed at Ilkemere (Fig. 8). Discussion The results of two herpetological surveys inthe SNP along parts of the eastern shore of Lake Turkana in northern Kenya are presented here. In addition to the records of 28 reptiles and six amphibians from these surveys, a June 2023 | Volume 17 | Number 1/2 | e324 Kirchhof et al. checklist of the herpetofauna in an extended area east of Lake Turkana from the Ethiopian border in the north, east to the town of North Horr, and south to Mount Kulal was presented at the 6" Congresso Nazionale della Societas Herpetologica Italica in Rome, Italy in 2006 (Ziliani et al 2006). Our surveys added seven species of reptiles and two amphibians that were not found during the 10 herpetological surveys conducted by Ziliani et al. (2006), despite their coverage of a larger range and spending more time. These nine species are: Holodactylus africanus, | Homopholis fasciata, — Philochortus rudolfensis, Psammophis cf. tanganicus, Ptychadena cf. schillukorum, Rhamphiophis rostratus, Sclerophrys xeros, Varanus albigularis, and Varanus niloticus. Thirteen reptile species and two amphibians recorded by Ziliani et al. (2006) in xeric habitats were not detected in our surveys. Most likely these taxa also find suitable habitat in SNP and are expected to occur there (Table 1). We did not consider the species that were recorded by Ziliani et al. (2006) in the very ecologically different, more mesic vegetation types, including the afromontane forest remnants at Mount Kulal (14 species, excluding Lygosoma afrum, currently considered a synonym of M. sundevallii and Psammophis semivariegatus, which does not exist and is likely Philothamnus semivariegatus; Table 1). Among the unique environmental features of the area for the local herpetofauna are (i) Lake Turkana, a permanent water source in this semi-desert, with a high pH and high salinity (for aquatic animals) and (11) the terrestrial shoreline of the lake with localized grass- dominated habitats. Our results show the lowest species diversity in those grasslands, a fact that was unexpected. However, the grasslands seem to be experiencing the highest impact from local livestock overgrazing. In addition, the prolonged droughts in the area likely affect the grasslands through changes in the flooding regime and seasonal lake-level fluctuations. Along the shore, the four species that were not recorded anywhere else can apparently withstand the rather inhospitable chemical conditions of the lake water. Those four species are: Crocodylus niloticus, except for one individual that was found a little further inland near one of our bushland transects; Varanus niloticus, a monitor lizard that lives close to water; Platyceps brevis, this species is not aquatic and was recorded in anthropogenic habitat between the stones of the wall of a building; and the toad Sclerophrys turkanae. Furthermore, although they were not recorded alive during our expeditions, three aquatic species of turtles and terrapins (Pelusios broadleyi, Pelomedusa subrufa [sic], now P. neumanni, and Trionyx triunguis) are restricted to the lake (Ziliani et al. 2006). These species are subject to fishing pressure, and they often end up as bycatch, are disturbed by fishermen, become entangled in nets, or take baited fishing hooks and drown (IUCN World Heritage Outlook 2020). The dry riverbeds also represent an important habitat, especially in such a dry area, mainly due to their high groundwater levels which are mandatory for the trees growing in the narrow riparian woodlands along the river. Consequently, we found most of the amphibian species in the riverbeds (Pitychadena nilotica, Amphib. Reptile Conserv. Ptychadena cf. schillukorum, Sclerophrys turkanae, and Tomopterna wambensis), as well as the more or less arboreal reptile species (Agama lionotus, Hemidactylus barbierii, Hemidactylus ruspolii, Homopholis fasciata, and Lygodactylus somalicus), although many of the latter also inhabited the shrubs in the bushland. Furthermore, the three individuals of Varanus albigularis were found exclusively along the riverbeds under loose bark of trees. The number of individuals we recorded 1s not acompletely accurate representation of abundance because not every individual encountered was caught and marked. As a result, on each second surveying day per transect there was a possibility of re-recording of individuals. In addition to the aforementioned aquatic and arboreal species, the recorded herpetofauna of the SNP comprises species typical of the semi-arid to arid savannas of East Africa. A few of the species, such as Bitis arietans, Heliobolus spekii, Hemidactylus platycephalus, — Homopholis _ fasciata, | Mochlus sundevallii, Rhamphiophis rostratus, R. rubropunctatus, Trachylepis quinquetaeniata, T. striata, T: varia, and Varanus albigularis, are widespread and generalist, also inhabiting moist and dry savanna areas. These species are generally distributed further to the south and enter parts of Central Africa. Furthermore, a few Palearctic taxa reach the Turkana area, such as Echis pyramidum and Stenodactylus sthenodactylus, as well as the largely Palearctic genus Eryx which is represented in SNP by E. colubrinus. Environmental Changes over Recent Decades Recently, the area around SNP has received more attention from biologists and conservationists than in the past (e.g., Cabeza et al. 2016; Conenna et al. 2019; Junqueira et al. 2021; Torrents-Ticé et al. 2021). The overall outcomes and impressions of these various studies are that throughout the past decades, the xeric areas in northern Kenya, including Lake Turkana and SNP, have been facing severe anthropogenic pressures. A 2 °C rise in minimum and maximum temperatures between 1967 and 2012 in the Turkana area has been reported (Avery 2012), as well as changes in the intensity and frequency of rainy seasons and increased duration and frequency of severe droughts (Junqueira et al. 2021). Ethnographic studies also report that the Daasanach people in the area have perceived increases in temperature and wind strength, drier and less fertile soils, less grass and increased water salinity (Junqueira et al. 2021). Photographic evidence from the 1960s shows that there was once a rich mammalian fauna in SNP, with giraffes, lions, and cheetahs, all of which are now locally extirpated (IUCN World Heritage Outlook 2020), and the local elders reported fertile and green pastures for their livestock in the past (Cabeza et al. 2016). Herders also report changes in the population trends of many of the mammals in the area (Torrents-Tico et al. 2021). Whether these changes are also affecting the herpetofauna of SNP is less clear, since most reptiles and amphibians are less targeted by the local population than the mammals. Nevertheless, changes in the lake water levels and increased anthropogenic pressures with negative impacts June 2023 | Volume 17 | Number 1/2 | e324 The Herpetofauna of Sibiloi National Park, Kenya on the vegetation cover (shade), insect abundance (reptile and amphibian prey), ground water levels, and land use practices (agro-pastoralism, fishing), are likely to have already impacted the diversity and abundance of reptile and amphibian species. Our results show that the majority of the herpetofauna of the SNP comprises desert and semi-desert taxa. Museum specimens are very rare for the eastern shore of Lake Turkana, which motivated us to collect voucher specimens of the local herpetofauna for the NMK collection to serve as a baseline for future generations of conservationists and biologists. The reptile and amphibian species recorded at Mount Kulal, which is about 100 km south of SNP and covered by afromontane forest remnants and mesophilous vegetation formations, show a herpetofauna community comprising species that typically occur in mesic savannas and _ forests further to the south and west, e.g., Bufo (Sclerophrys) gutturalis, Crotaphopeltis hotamboeia, Dispholidus typus, Gerrhosaurus flavigularis, and Philothamus semivariegatus [sic] (Philothamnus semivariegatus) (Ziliani et al. 2006). These species seem to represent relict populations for the area, and they were likely to be more widespread under the less severe environmental conditions which were reportedly still prevalent in the Turkana region less than 50 years ago (Cabeza et al. 2016; Junqueira et al. 2021; Torrents-Tico et al. 2021). The opportunistically recorded field body temperatures of some of the species (see Species Accounts) were not exceptionally high, so we assume that at least those taxa were able to effectively thermoregulate in the SNP despite the increased environmental temperatures (Avery 2012). Conclusions The SNP harbors a high diversity of reptiles, and also a decent amount of amphibian taxa for a xeric environment, and many of these taxa are poorly studied. During two surveys, a number of species were found for the first time in SNP, although none of them were unexpected based on their known ecology and distribution ranges. There are probably some more species to discover in the area, and splitting taxa based on molecular analyses might further extend the species list. Based on the present survey and the expeditions by Ziliani et al. (2006), the currently known reptile and amphibian fauna of the SNP comprises A9 species, including eight amphibians and 41 reptiles (three freshwater turtles, one crocodile, 25 lizards, and 12 snakes). The herpetofauna includes species typical of the semi- arid to arid savannas of East Africa (including arboreal taxa), a few taxa with more Palearctic distributions, a few widespread and generalist taxa that also inhabit more humid areas, and the aquatic species dependent on Lake Turkana. The latter group comprises four reptile species listed on CITES App. II and two reptile species listed on the IUCN Red List, highlighting the importance of the lake for the Reptilia. These species are also the most likely group to be affected if the water resources, including the amount of available water and the chemistry of the lake, are negatively impacted by climate change Amphib. Reptile Conserv. and the Gilgel Gibe III Dam in Ethiopia. In addition, the habitats associated with the ephemeral rivers that have high groundwater levels and riparian woodlands/forests, which are home to most amphibian species and many reptile species, will be heavily affected if water levels drop. The grassy habitats along the lake shore seem to be impacted by overgrazing and now harbor fewer reptile and amphibian species than expected. Overall, the herpetofauna of this area includes a number of CITES and IUCN Red List listed species, including endemics, that warrant protection and conservation measures to prevent further defaunation. Acknowledgments.—We thank the National Commission for Science, Technology, and Innovation (NACOSTI/P/16/21446/14491) and Kenya Wildlife Service (KWS/BRM/5001) for granting access to the area, and the Turkana Basin Institute (TBI) for all their logistic support. This study has received funding from the Nordenskidld Expedition fund (granted to Mar Cabeza, University of Helsinki), and funds for the promotion and support of young researchers from the Museum fir Naturkunde Berlin (Germany) (granted to Sebastian Kirchhof). The authors wish to thank Mikael Fortelius and Mar Cabeza (both at University of Helsinki) for offering the opportunity to join the two expeditions to SNP, and Shooro Claudia Goosh, Lawrence Losogo Bosco, Irene Conenna, the Helsinki team, and the TBI team for their field assistance, team spirit, general help, and company. Mar Cabeza also added helpful comments to the manuscript. Special thanks go to Eli Greenbaum for his suggestions which improved the article. Literature Cited Avery S. 2012. Lake Turkana and the Lower Omo: Hydrological Impacts of Major Dam and Irrigation Developments. Report. African Studies Centre, University of Oxford, Oxford, United Kingdom. 239 p. BirdLife International. 2022. 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Revision of the African snakes of the genera Dromophis and Psammophis. Bulletin of the Museum of Comparative Zoology 87(1): 1-69. Mbaluka JK, Brown FH. 2016. Vegetation of the Koobi Fora Region northeast of Lake Turkana, Marsabit County, northern Kenya. Journal of East African Natural History 105(1): 21-50. Nago SGA, Grell O, Sinsin B, R6del MO. 2006. The amphibian fauna of the Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93- 108. Olson DM, Dinerstein E, Wikramanayake ED, Burgess ND, Powell GV, Underwood EC, D’ Amicao JA, Itoua Amphib. Reptile Conserv. I, Strand HE, Kassem KR, et al. 2001. Terrestrial ecoregions of the world: a new map of life on Earth. A new global map of terrestrial ecoregions provides an innovative tool for conserving biodiversity. BioScience 51: 933-938. Sindaco R, Razzetti E, Ziliani U, Wasonga V, Carugati C, Fasola M. 2007. A new species of Hemidactylus from Lake Turkana, northern Kenya (Squamata: Gekkonidae). Acta Herpetologica 2: 37-48. Spawls S, Howell K, Hinkel H, Menegon M. 2018. Field Guide to East African Reptiles. Bloomsbury Publishing, London, United Kingdom. 624 p. Spawls S, Wasonga V, Drewes RC. 2019. The Amphibians of Kenya. Stephen Spawls (privately published). 55 p. Tolley KA, Alexander GJ, Branch WR, Bowles P, Maritz B. 2016. Conservation status and threats for African reptiles. Biological Conservation 204: 63-71. Thorsell J. 2003. World Heritage Nomination-IUCN Technical Evaluation. ulissat Icefjord (Denmark) ID No: 1149. IUCN, Gland, Switzerland. 5 p. Torrents-Tico M, Fernandez-Llamazares A Burgas D, Cabeza M. 2021. Convergences and divergences between scientific and indigenous and local knowledge contribute to inform carnivore conservation. Ambio 50(5): 990—1,002. Uetz P, Freed P, Aguilar R, HoSek J. (Editors). 2022. The Reptile Database. Available: http://www. reptile- database.org [Accessed: 1 April 2022]. UNEP-WCMC, IUCN. 2022. Protected Planet: The World Database on Protected Areas (WDPA) and World Database on Other Effective Area-based Conservation Measures (WD-OECM). UNEP- WCMC, Cambridge, United Kingdom and IUCN, Gland, Switzerland. Available: http://www. protectedplanet.net [Accessed: 10 July 2022]. Weinell JL, Branch WR, Colston TJ, Jackman TR, Kuhn A, Conradie W, Bauer AM. 2019. A species-level phylogeny of Trachylepis (Scincidae: Mabuyinae) provides insight into their reproductive mode evolution. Molecular Phylogenetics and Evolution 136: 183-195. Yuretich RF, Cerling TE. 1983. Hydrogeochemistry of Lake Turkana, Kenya: mass balance and mineral reactions in an alkaline lake. Geochimica et Cosmochimica Acta 47(6): 1,099-1,109. Ziliani U, Sindaco R, Razzetti E, Wasonga V, Modry D, Necas P, Carugati C, Fasola M. 2006. The Herpetofauna of the Eastern Side of the Lake Turkana (Northern Kenya). Pp. 192-193 In: Riassunti del 6° Congresso Nazionale della Societas Herpetologica Italica (Roma, 27.IX-1.X.2006). Editors, Bologna MA, Capula M, Carpaneto GM, Luiselli L, Marangoni C, Venchi A. Stiligrafica, Rome, Italy. 238 p. 9 June 2023 | Volume 17 | Number 1/2 | e324 Amphib. Reptile Conserv. The Herpetofauna of Sibiloi National Park, Kenya Sebastian Kirchhof is a research associate at New York University Abu Dhabi working in close collaboration with the Museum fiir Naturkunde Berlin, Germany, where he received his Ph.D. Sebastian 1s a trained ecologist, nature conservationist, and biologist, currently focusing on genomic adaptation to extreme conditions in amphibians and reptiles by combining genomics, transcriptomics, niche modeling, population genomics, and systematics. He is interested in the evolutionary processes responsible for the diversification of Earth’s biota and genomic architecture, and how they affect speciation rates. This knowledge is one of the fundamental pillars for the protection of the Earth’s biodiversity. Victor Wasonga is a trained conservation biologist and a research scientist based at the National Museums of Kenya. Victor’s current research involves using herpetofauna to understand phylogeny, taxonomy, ecology, distribution, landuse, ecosystem dynamics, and livelihoods. He has authored many publications and described a number of species. Victor is currently serving as a Co-Chair of the East African Amphibian Specialist Group within IUCN’s Species Survival Commission. Tomas Mazuch is a Czech amateur herpetologist. Since his teenage years, Toma’ has dedicated his life to the breeding of amphibians, reptiles, and invertebrates. During his studies of Veterinary Sciences (which are not finished yet), he began to devote his research to herpetology and the parasitology of reptiles. His research focuses on the taxonomy, systematics, and biogeography of amphibians and reptiles of the Horn of Africa (mainly Somalia and Ethiopia). His main subjects of study are geckos of the genus Hemidactylus from Eastern Africa. Tomas is also interested in the taxonomy of scorpions and succulent plants from North-Eastern Africa. He has authored or co- authored about 30 peer-reviewed papers and books on parasitology, the systematics of scorpions, plants, and herpetology, including the book Amphibians and Reptiles of Somaliland and Eastern Ethiopia, Based on Two Field Trips in 2010/2011 in 2013. He has co-authored the descriptions of 11 reptile, six scorpion and two plant species. He currently runs long-term projects and field studies in Somaliland. Stephen Spawls is an independent herpetologist who has worked extensively in Africa, in Kenya, Egypt, Ghana, Botswana, and Ethiopia. His publications include A Field Guide to the Reptiles of East Africa and a book on Africa’s dangerous snakes. He lives in Norwich, United Kingdom. Patrick Kinyatta Malonza has been a herpetologist at the National Museums of Kenya- Nairobi since 1996, where he leads a team of other herpetologists and enthusiasts to promote reptile and amphibian conservation in Kenya. Patrick works on the taxonomy, community ecology, and conservation of reptiles and amphibians. He has authored or co-authored over 35 publications. His key interest is understanding species and habitat relationships, species descriptions, and their biogeographical associations. 18 June 2023 | Volume 17 | Number 1/2 | e324 Official journal website: amphibian-reptile-conservation.org Amphibian & Reptile Conservation 17(1 & 2) [General Section]: 19-56 (e325). Contributions to the herpetofauna of the Angolan Okavango- Cuando-Zambezi river drainages. Part 3: Amphibians 1.2.3.* Werner Conradie, 2?Chad Keates, *°°Luke Verburgt, *7%°°"Ninda L. Baptista, and *'2James Harvey 'Port Elizabeth Museum, Beach Road, Humewood, Port Elizabeth 6013, SOUTH AFRICA *Department of Nature Conservation Management, Natural Resource Science and Management Cluster, Faculty of Science, George Campus, Nelson Mandela University, George, SOUTH AFRICA 3National Geographic Okavango Wilderness Project, Wild Bird Trust, SOUTH AFRICA *South African Institute for Aquatic Biodiversity (SATAB), Makhanda, SOUTH AFRICA °*Enviro-Insight CC, Unit 8 Oppidraai Office Park, Pretoria 0050, SOUTH AFRICA ‘Department of Zoology and Entomology, University of Pretoria, Pretoria, 0001, SOUTH AFRICA ‘Instituto Superior de Ciéncias da Educagado da Huila (ISCED), Rua Sarmento Rodrigues, Lubango, ANGOLA ®CIBIO/InBio, Centro de Investigacdo em Biodiversidade e Recursos Genéticos, Laboratorio Associado, Universidade do Porto, Campus Agrario de Vairdo, Rua Padre Armando Quintas, 4485-661 Vairdo, PORTUGAL °Departamento de Biologia, Faculdade de Ciéncias, Universidade do Porto, 4169-007, Porto, PORTUGAL '°BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Campus de Vairdo, 4485-661 Vairdo, PORTUGAL ''Museum fiir Naturkunde—Leibniz Institute for Evolution and Biodiversity Science, Invalidenstr. 43, 10115 Berlin, GERMANY "Harvey Ecological, 41 Devonshire Avenue, Howick, 3290, SOUTH AFRICA Abstract.—This article is the third and final installment of the herpetofaunal results obtained from a series of rapid biodiversity surveys of the upper Cuito, Cubango, Cuando, Zambezi, and Kwanza River basins in Angola. The amphibian survey results are presented along with an updated checklist of the historical and current records of amphibians from the southeastern region of Angola. A total of 1,114 new amphibian records were documented, comprising 37 species, bringing the total number of recognized amphibian species in this region to 49. These surveys documented two new country records (Hyperolius cf. inyangae and Kassinula wittei) and at least two candidate new species, and elevated Amnirana adiscifera stat. nov. (which now encompasses the western green form formerly regarded as A. darlingi). Finally, updated distribution maps for all of Angola are provided for all the species encountered within the study region. Keywords. Africa, Cuanavale, Cuito, Okavango, headwaters, frogs Resumo.—Este trabalho é a terceira e ultima parte dos resultados obtidos de uma série de levantamentos rapidos de biodiversidade realizados nas bacias dos rios Cuito, Cubango, Cuando, Zambeze e Kwanza em Angola. Aqui apresentamos uma lista atualizada de registos anfibios do sudeste de Angola, consistindo em registos historicos e actuais. Ao todo foram obtidos 1,114 novos registos, relativos a 37 espécies, elevando o numero total de especies de anfibios reconhecidas nesta regiao para 49. Duas especies foram registadas no pais pela primeira vez (Hyperolius cf. inyangae, Kassinula wittei). Os nossos resultados sugerem a existéncia de pelo menos duas potenciais especies novas, e elevaram o estatuto de Amnirana adiscifera stat. nov. para abranger a forma verde ocidental de A. darlingi. Por fim, produzimos mapas de distribuigao actualizados para todas as especies encontradas neste estudo para 0 pais inteiro. Palavras-chave. Africa, Cuanavale, Cuito, Okavango, nascentes, sapos Citation: Conradie W, Keates C, Verburgt L, Baptista NL, Harvey J. 2023. Contributions to the herpetofauna of the Angolan Okavango-Cuando- Zambezi River drainages. Part 3: Amphibians. Amphibian & Reptile Conservation 17(1 & 2) [General Section]: 19-56 (e325). Copyright: © 2023 Conradie et al. This is an open access article distributed under the terms of the Creative Commons Attribution License [Attribution 4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org. Accepted: 13 April 2023; Published: 4 August 2023 Introduction In recent years, knowledge on the Angolan herpetofauna has increased dramatically (Marques et al. 2018; Baptista et al. 2019; Branch et al. 2019a). Although reptiles have been the main focus (Conradie et al. 2012a, 2020a, 2021, 2022a,b,c; Stanley et al. 2016; Ceriaco et al. 2020a,b,c; Marques et al. 2019a,b, 2020, 2022a,b, 2023; Branch et al. 2019a,b, 2021; Hallermann et al. 2020; Lobon- Rovira et al. 2021, 2022; Parrinha et al. 2021; Wagner et al. 2021), several amphibian-focused studies have been published (Conradie et al. 2012b, 2013, 2020b; Ernst et al. 2014, 2015; Conradie and Bills 2016; Ceriaco et al. 2018, 2021; Nielsen et al. 2020; Baptista et al. 2021). Correspondence. *werner@bayworld.co.za, wernerconradie@gmail.com Amphib. Reptile Conserv. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola From these studies, only six new species of amphibians have been described for Angola since 2012, compared to 35 new species of reptiles (29 lizards and six snakes). Recent syntheses on the amphibians of Angola documented between 111 and 117 anuran species within the country (Marques et al. 2018; Baptista et al. 2019), although the taxonomic status and presence within Angola remains uncertain for several of these frogs. Approximately 22 species are known only from the type description, singleton records, or incorrect species assignments which no longer apply to the Angolan material (Marques et al. 2018; Baptista et al. 2019). A large proportion of historical type material was lost or destroyed during the Natural History Museum Lisbon fire (Marques et al. 2018), complicating matters further, thus new topotypic material is needed to validate the taxonomical status of various species (Baptista et al. 2019). Over the last few years, additional amphibian species have been either recorded for the first time from Angola (Conradie et al. 2020b; Ernst et al. 2020) or described as new species (Ceriaco et al. 2018, 2021; Nielsen et al. 2020; Baptista et al. 2021), raising the number of amphibian species known from Angola by at least 13. As more work is done in the country and remote regions are surveyed, a trend similar to reptiles is expected, with the predicted discovery and description of multiple new species. Southeastern Angola remains amongst the most poorly known regions in Africa (Conradie et al. 2016; Marques et al. 2018; Baptista et al. 2019). Historical amphibian records from this region included only opportunistic collections, most of which were restricted to the western tributaries of the Cubango River basin (Bocage 1895; Monard 1937), and the eastern and northern sections of Moxico Province (Laurent 1964; Monard 1937; Mertens 1938; Ruas 1996, 2002), with only a few records from the Cuito and Cuando river basins (Angel 1924). Recently, based on the outcome of several biodiversity surveys (Brooks 2012, 2013; NGOWP 2017) to document the biodiversity of the Angolan catchment of the Okavango River basin, which comprises the Cubango, Cuito, and Cuando rivers, an updated herpetofauna checklist was compiled (Conradie et al. 2016). A total of 34 species of amphibians were recorded, four of which were new for Angola (Ptychadena mossambica, Sclerophrys poweri, Xenopus muelleri, and X. poweri). The present work serves as the third and final installment of a series of papers documenting the herpetofauna of southeastern Angola, based on surveys performed for the National Geographic Okavango Wilderness Project. The first two installments focused on snakes (Conradie et al. 2021) and on lizards, chelonians, and crocodiles (Conradie et al. 2022c), while this article is restricted to amphibians. The overarching aim of these articles is to document and quantify the herpetofauna of the region, and to contribute to the knowledge of its Amphib. Reptile Conserv. conservation importance in both regional and national contexts. Materials and Methods For this study, we present amphibian material and associated data collected during five National Geographic Okavango Wilderness Project (NGOWP) expeditions to south-eastern Angola from 2016 to 2019. See Conradie et al. (2021) for more details on these surveys, a description of the study area, sampling techniques (trapping and visual encounter surveys), and species mapping. Below are some specific methods pertaining to this article. Identification and morphology. Upon completion of the fieldwork component of the study, species were identified based on external morphology, using relevant field guides or published identification keys (Poynton and Broadley 1985a,b, 1987, 1988; Channing 2001; Du Preez and Carruthers 2017; Channing and Rodel 2019) and orginal type descriptions when needed, and through direct comparisons with material housed in the Port Elizabeth Museum (PEM). Identifications of tadpoles were problematic, as the tadpoles of many of the species collected as adults remain undescribed (see Channing et al. 2012). We tentatively assigned tadpoles based on morphotypes and locality data to the known species until their true identities can be confirmed by genetic analysis. In certain cases, a 16S rRNA barcoding approach was employed to aid in the identification of tadpoles and adults. Laboratory and sequencing protocols followed Conradie et al. (2020b). The Basic Local Alignment Search Tool (BLAST; Altschul et al. 1990) was used to compare our material to the GenBank (https://www.ncbi. nlm.nih.gov/genbank/; Benson et al. 2013) repository and our unpublished Angolan dataset. Taxonomy follows Frost (2023) and was updated as needed. Common names follow Channing (2001), Du Preez and Carruthers (2017), and Marques et al. (2018). The snout-urostyle length (SUL, measured from the tip of the snout to the posterior tip of the urostyle) of adult Specimens and the total length of tadpoles were measured to the nearest 0.1 mm, using a digital calliper. All specimens were examined using a Nikon SMZ1270 binocular stereo microscope. For adult frogs, the following information was documented: skin texture, position and number of hand/ feet tubercles, webbing condition, and coloration. The webbing formula follows the scheme provided by Rodel (2000). Adult specimens were sexed by confirming the presence/absence of eggs, a gular flap or darkened throats, and nuptial pads. As needed, a small ventral incision was made to look for gonads or testes. No attempt was made to examine stomach contents (which will be the focus of another study). Additionally, the labial tooth rows and oral papillae condition were recorded for tadpoles. Tadpoles are catalogued as ‘lots,’ with specimens from the same locality and collecting event grouped together. August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. ene & Ne : a? = Fig. 1. Adult male Arthroleptis stenodactylus from Lungwebungu River. Photo by Werner Conradie. Results We documented a total of 1,114 (1,011 adult frogs and 103 tadpole lots) individual amphibian records from ~189 unique localities in southeastern Angola, particularly around the source lakes of the Cuito, Cuanavale, Cuando, and Quembo rivers. A total of 36 species of amphibians (comprising eight families and 20 genera, all from the order Anura) were recorded during this study (Table 1). Information is also reported for one additional species (Hyperolius quinquevittatus) that was not collected from within the core study area as previously defined. Updated species occurrence maps are provided for each of the 37 species, reflecting all known localities in Angola (Maps 1-37). For the mapping exercise, a total of 2,507 unique records were collated: 1,062 historical records from Marques et al. (2018), 522 additional literature records, 149 citizen science platform records, 296 records from other sources (GBIF ~ https://www. gbif.org/, unpublished records of the PEM, SATAB, and ZMB collections), and 925 unpublished records from our surveys. This mapping exercise increased the number of new or previously undocumented amphibian records in this study by 58%. Below we provide a checklist of the amphibians found during these surveys, with each entry including a list of the material examined and comments on identification, habitat, distribution, taxonomy, and natural history notes. Material not collected by the core team or examined for this paper are referred to under ‘Additional material.’ New data used to compile distribution maps can be found in Supplementary Table 1 at: https://doi.org/10.6084/m9 figshare.23544306.v1. Abbreviation used: asl — above sea level. Museum and col- lectors’ codes used: INBAC — Instituto Nacional de Gestaéo Ambiental; PEM — Port Elizabeth Museum; ZMB — Museum fur Naturkunde Berlin (Zoological Collections); SAIAB — South African Institute for Aquatic Biodiveristy; P — Pedro vaz Pinto, WC — Werner Conradie. Amphibia Arthroleptidae Amphib. Reptile Conserv. 21 Elevation (m) 14 -16 -18 18 20 22 24 Map 1. Distribution of Arthroleptis stenodactylus in Angola. Historical records (Marques et al. 2018) are indicated by white dots, while new records are indicated by black dots. Axis values are in degrees (°). Purple polygon — Okavango River basin, Blue polygon — Cuando River basin, Brown polygon — Zambezi River basin. Arthroleptis stenodactylus Pfeffer, 1893 Common Squeaker (Fig. 1; Map 1) Material (43 specimens): PEM A12495, 4 km upstream from Cuanavale River source lake, -13.05084° 18.89726°, 1,395 m asl; PEM A12501—4, river crossing before Samboano village, -12.30700° 18.62350°, 1,397 m asl; PEM A12526, Munhango village, -12.16310° 18.55430°, 1,421 m asl; PEM A12527, Protea stop en route to Cuito River source, -12.30040° 18.62070°, 1,429 m asl; PEM A12528—36, INBAC: (no number x4), Cuito River source lake, -12.68935° 18.36012°, 1,431 m asl; PEM A12585, 10 km north of Cuemba village, -12.03481° 18.04869°, 1,372 m asl; PEM A13815, en route to Lungwebungu River, -12.25034° 18.63742°, 1,506 m asl; PEM A12616-7, en route road to Cuito River source, -12.55152° 18.41434° 1,507 m asl; PEM A12618, en route road to Cuito River source, -12.25050° 18.63730°, 1,519 m asl; PEM A12620, drive to Cuanavale River camp from Samanunga village, -13.03803° 18.82977°, 1,623 m asl; PEM A12644, Cuanavale River source lake, -13.18067° 18.92172°, 1,340 m asl; PEM A12647, stop on road to Cuito River source, -12.50584° 18.41382°, 1,556 m asl; PEM A12648-9, camp | en route to Cuito River source, -12.35920° 18.56280°, 1,510 m asl; PEM A12729, Cuando River source, -13.00346° 19.12751°, 1,346 m asl; PEM A12803-4, Lake Tchanssengwe, -12.41402° 18.64418°, 1,415 m asl; PEM A12843-—50, INBAC: (no number x2), Quembo River source lake, -13.13624° 19.04591°, 1,411 m asl. Additional records: P2-276 (photograph and tissue sample), wetland near old quarry east of Quemba, -12.16960° 18.22965°, 1,353 m asl. Description: Medium sized Arthroleptis, stocky build; rounded snout; tympanum clearly visible; well- developed inner metatarsal tubercle; no webbing; toe tips not dilated. Dorsum uniformly beige to pink, with scattered white speckles; no darker hourglass pattern observed on dorsum; yellow vertebral stripe either August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola Table 1. Records of amphibians for the Angolan Okavango-Cuando-Zambezi river basins. ? = not recorded from the core study area, but expected to occur based on peripheral records. Species ARTHROLEPTIDAE Arthroleptis stenodactylus Pfeffer, 1893 Arthroleptis xenochirus Boulenger, 1905 Leptopelis anchietae (Bocage, 1873) Leptopelis bocagii (Ginther, 1865) Leptopelis sp. BREVICIPITIDAE Breviceps ombelanonga Nielsen, Conradie, Ceriaco, Bauer, Heinicke, Stanley, and Blackburn, 2020 BUFONIDAE Mertensophryne melanopleura (Schmidt and Inger, 1959) Poyntonophrynus kavangensis (Poynton and Broadley, 1988) Schismaderma carens (Smith, 1848) Sclerophrys funerea (Bocage, 1866) Sclerophrys gutturalis (Power, 1927) Sclerophrys pusilla (Mertens, 1937) Sclerophrys poweri (Hewitt, 1935) Sclerophrys lemairii (Boulenger, 1900) HEMISOTIDAE Hemisus guineensis Cope, 1865 HYPEROLITDAE Hyperolius benguellensis (Bocage, 1893) Hyperolius cinereus Monard, 1937 Hyperolius cf. invangae Channing, 2013 Hyperolius nasutus Gunther, 1865 FHyperolius parallelus Gunther, 1858 Hyperolius raymondi Conradie, Branch, and Tolley, 2013 Hyperolius quinquevittatus Bocage, 1866 Hyperolius aff. bocagei Steindachner, 1867 Amphib. Reptile Conserv. Okavango River Basin ~< x x KK XK Cuando River Basin 22 x x Zambezi River Basin x x KK Source of records This study; Ruas 1996 This study; Ruas 1996 This study; Monard 1937; Conradie et al. 2016 This study This study This study; Bocage 1895; Monard 1937; Ruas 1996 Ruas 1996 Ruas 1996 Laurent 1964 Monard 1937; Laurent 1964; Ruas 1996; Conradie et al. 2016 This study; Monard 1937; Ruas 1996; Conradie et al. 2016 This study This study; Conradie et al. 2016 Ruas 1996; Conradie et al. 2016 This study; Monard 1937; Laurent 1964 This study; Conradie et al. 2016 This study; Conradie et al. 2016 This study This study; Monard 1937; Conradie et al. 2016 This study; Conradie et al. 2016 This study This study This study August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. Table 1 (continued). Records of amphibians for the Angolan Okavango-Cuando-Zambezi river basins. ? = not recorded from the core study area, but expected to occur based on peripheral records. Species Kassina kuvangensis (Monard, 1937) Kassina senegalensis (Duméril and Bibron, 1841) Kassinula wittei Laurent, 1940 MICROHYLIDAE Phrynomantis affinis Boulenger, 1901 PHRYNOBATRACHIDAE Phrynobatrachus mababiensis FitzSimons, 1932 complex Phrynobatrachus natalensis (Smith, 1849) Phrynobatrachus parvulus (Boulenger, 1905) PIPIDAE Xenopus muelleri (Peters, 1844) Xenopus petersii Bocage, 1895 Xenopus poweri Hewitt, 1927 PTYCHADENIDAE Hildebrandtia ornatissima (Bocage, 1879) Ptychadena anchietae (Bocage, 1868) Ptychadena bunoderma (Boulenger, 1907) Ptychadena grandisonae Laurent, 1954 Ptychadena guibei (Laurent, 1954) Ptychadena keilingi (Monard, 1937) Ptychadena nilotica (Seetzen, 1855) Ptychadena mossambica (Peters, 1854) Ptychadena oxyrhynchus (Smith, 1849) Ptychadena porosissima (Steindachner, 1867) Ptychadena subpunctata (Bocage, 1866) Ptychadena taenioscelis Laurent, 1954 Ptychadena upembae (Schmidt and Inger, 1959) Amphib. Reptile Conserv. Okavango River Basin Cuando River Basin 23 x Zambezi River Basin ~< x x KM Source of records This study; Monard 1933, 1937; Conradie et al. 2016 This study; Monard 1933, 1937; Conradie et al. 2016 This study; Conradie et al. 2021 Ruas 1996 This study; Conradie et al. 2016 This study; Monard 1937; Ruas 1996; Conradie et al. 2016 Conradie et al. 2016 Conradie et al. 2016 This study; Monard 1937; Ruas 1996; Conradie et al. 2016 This study; Conradie et al. 2016 Bocage 1895; Monard 1937a; Ruas 1996 Ruas 1996 This study This study, Ruas 1996 Ruas 1996 This study; Ruas 1996 Ruas 1996; Conradie et al. 2016 Conradie et al. 2016 This study; Monard 1937a; Conradie et al. 2016 This study This study; Ruas 1996; Conradie et al. 2016 This study; Ruas 1996; Conradie et al. 2016 This study; Ruas 1996; Conradie et al. 2016 (as P. guebei) August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola Table 1 (continued). Records of amphibians for the Angolan Okavango-Cuando-Zambezi river basins. ? = not recorded from the core study area, but expected to occur based on peripheral records. ’ Okavango Species . ; River Basin Ptychadena uzungwensis (Loveridge, 1932) x PYXICEPHALIDAE Amietia angolensis (Bocage, 1866) x Tomopterna tuberculosa (Boulenger, 1882) x RANIDAE Amnirana adiscifera (Schmidt and Inger, 1959) stat. nov. x Amnirana lemairei (De Witte, 1921) Species total: 51 40 absent (n = 23) or present (n = 12); faint dark facial mask from tip of snout to the eye, continuous to the arm; ventrum immaculate. Breeding males with dark throats; minute spines on lower back; elongated 3” finger, with spines on the inner side and to the tip. Adult females (n = 11) varied from 32.6—37.7 (34.6 average) mm SUL (largest female: PEM A12846); adult males (n = 4) varied from 19.5—23.6 (21.5) mm (largest male: PEM A12804). Habitat and natural history notes: All specimens were collected in miombo woodland. Gravid females were collected in October/November. Males were heard calling on overcast days and evenings after rain. In February/March, juveniles and subadults were abundant, but no adults were collected or heard calling. Comments: Historically, this species was only known from three records from eastern Angola (Marques et al. 2018). Our new material shows that this species is more Fig. 2. Adult male Arthroleptis xenochirus from Muhango town. Photo by Werner Conradie. Amphib. Reptile Conserv. Cuando ; Zambezi River . ; Source of records . River Basin Basin This study; Ruas 1996; Conradie et x x al. 2016 This study; Ruas 1996; Conradie et x al. 2016 This study; Bocage 1895; Monard xX 1937; Ruas 1996 This study; Ruas 1996; Conradie et x x al. 2016 x Ruas 1996 31 42 common in eastern Angola than previously recognized. This was expected as it 1s widespread to the east of the Zambian border (Poynton and Broadley 1985a, 1991; Channing 2001). Additionally, these are the first confirmed records from the Okavango River basin. Studies have shown that A. stenodactylus comprises two distinct ecomorphs that occur in different habitats. One form, to which our material belongs, prefers drier savanna, while the other form is known from montane forests (Loveridge 1953; Pickersgill 2007; Bittencourt-Silva et al. 2020). Arthroleptis xenochirus Boulenger, 1905 Plain Squeaker (Fig. 2; Map 2) Material (8 specimens): PEM A12505-7, river crossing before Samboano Village, -12.30700° 18.62350°, 1,398 m asl; PEM A12910-1, Muhango village, -12.16067° 18.55042°, 1,430 m asl; PEM A14696-7, Lungwebungu Elevation (m) 3 oO o Map 2. Distribution of Arthroleptis xenochirus in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. River camp, -12.58439° 18.66748°, 1,297 m asl; PEM A14777, Quembo River bridge camp, -13.52746° 19.2806°, 1,241 m asl. Additional material: P2-272 (photograph and tissue sample), wetland near old quarry east of Quemba, -12.16960° 18.22965°, 1,353 m asl. Description: A small species of Arthroleptis, slender body; pointed snout; small tympanum, not always clearly visible; well-developed inner metatarsal tubercle; no webbing; slightly dilated toe tips. Dorsal color varies from beige to brown, with some red infusion; dark facial mask from snout to eye, continuing to just above arm: white markings on grey colored jawline; all specimens have a darkened hour-glass pattern on the back; ventrum immaculate. Breeding males with dark throats; elongated 3 finger, with no spines. Single adult female measured 20.9 mm (PEM A14777); adult males (n = 4) varied from 19.5—23.6 (21.5) mm (largest male: PEM A14697). Habitat and natural history notes: All specimens were collected in miombo woodland. Males were heard calling in October/November. Gravid female collected in November, while only juveniles and subadults were collected in February/March. Comments: Known from central and northern Angola (Marques et al. 2018). The record from lower Quembo River represents the southernmost record for the species, and the first record associated with the Okavango River basin. Leptopelis anchietae (Bocage, 1873) Anchieta’s Tree Frog (Map 3) Material (1 tadpole lot): PEM A14174 (five tadpoles), Cubango River campsite below rapids, west of Fundo village, -13.04260° 16.37476°, 1,559 m asl. Description: Elongated tadpoles; 20.8—30.6 mm total length, with tail 2.6-3.1 times the body length. Body is dark brown to black, ventrum with scattered light golden spots; lateral tail muscle dark brown to black, with two lighter bands from body to tail tip; posterior half of tail black. The labial tooth row formula (LTRF) 1s 3(2-3)/3(1); jaw sheaths are heavily keratinized; anterior part of mouth free of elongated marginal papillae. Habitat and natural history notes: Tadpoles were collected in a flooded grassland next to the main river. Comments: f Elevation (m) Map 3. Distribution of Leptopelis anchietae in Angola. Amphib. Reptile Conserv. Identification of tadpoles was confirmed by 16S rRNA barcoding (N. Baptista, unpub. data) and compared to the description in Channing et al. (2012). This Angolan endemic species occurs mostly in the highlands of central and western Angola (Becker et al., in prep.), and many of the old historical records (e.g., Marques et al. 2018) are based on incorrect identifications or erroneous locality data (Pedro vaz Pinto, pers. comm. ). Leptopelis bocagii (Gunther, 1865) Bocage’s Tree Frog (Figs. 3—4; Map 4) Material (46 specimens): PEM A12701-10, Lungwebungu River camp bridge crossing, -12.58347° 18.66598°, 1,304 m asl; PEM A12742-6, Cuando Fig. 3. Adult male Leptopelis bocagii (brown form) from Cuando River source. Photo by James Harvey. ee ay “ £ ca ie ) at a, ff A... oc.) a as — oe. FS) a oe: es : = _ * Med a . Fig. 4. Adult male Leptopelis bocagii (green form) from Cuanavale River side tributary source. Photo by Luke Verburet. Elevation (m) 3 fo) oO 12 14 16 18 20 22 24 Map 4. Distribution of Leptopelis bocagii in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola River source, -13.00346° 19.12751°, 1,346 m asl; PEM A12772-4, INBAC: WC-4634, Cuanavale River side tributary source, -13.07452° 18.88345°, 1,385 m asl: PEM A12873-80, INBAC: WC-4669 and 4672, Quembo River source lake, -13.13624° 19.04591°, 1,411 m asl; PEM A14688-9, Lungwebungu River camp, massambas on left side of river, -12.58276° 18.66556°, 1,295 m asl; PEMA14701—4, INBAC: WC-6763, Lungwebungu River camp, -12.58439° 18.66748°, 1,297 m asl; PEM A14723, PEM A14742, Quembo River bridge camp, -13.52746° 19.28060°, 1,241 m asl; PEM A14741, Quembo River bridge camp, trap 3, -13.52778° 19.27455°, 1,256 m asl; PEM A14755-6, PEM A14758-9, INBAC: WC-6994, Quembo River right side tributary (Micongo River) past village, -13.51877° 19.28487°, 1,248 m asl; PEM A14823, camp at side tributary (Luandai River) of the Luanguinga River, -13.70885° 21.26234°, 1,116 m asl; PEM A14891, Luvu River camp, -13.71200° 21.83538°, 1,082 m asl. Description: Large terrestrial tree frog; broad rounded head; large protruding eyes; tympanum clearly visible; large well-developed inner metatarsal tubercle; no webbing or expanded toe tips. Dorsum varied from green to brown, with dark horseshoe pattern on the back, and dark interorbital bar; dark facial mask from snout to eye, continuing to above arm; sides of body with scattered black spots or continuous black bar between limbs; scattered white speckles on dorsum; ventrum immaculate. Breeding males with dark throat and weakly developed pectoral glands. Adult females (n = 12) varied from 41.5-67.8 (57.9) mm (largest female: PEM A12746); adult males (n = 34) varied from 43.0—56.8 (48.6) mm (largest male: INBAC: WC-4669). Habitat and natural history notes: Collected along the sandy margins of source lakes or rivers associated with miombo woodland. Frequently encountered in the clearings of agricultural fields near water sources. Males were found calling from the ground. Comment. These are the first records from eastern Angola, bridging the distribution gap between western Angola and Zambia (Broadley 1971; Marques et al. 2018; Baptista et al. 2019). Leptopelis sp. Unidentified Tree Frog (Figs. 5-6; Map 5) Material (34 specimens, 2 tadpole lots): PEM A12801-—2, Cuanavale River source lake, -13.08934° 18.89485°, 1,359 m asl; PEM A12882-8, Quembo River source lake, -13.13624° 19.04591°, 1,396 m asl; PEM A12794—5, INBAC: WC-4685 and no number, Cuanavale River source lake, -13.09442° 18.89372°, 1,368 m asl; PEM A12747-51, INBAC: WC-4754 and no number, Cuando River source, -13.00346° 19.12751°, 1,353 m asl; PEM A14118 (tadpoles), Calua River source, 6 km SE of Cuito River source, -12.73675° 18.39310°, 1,446 m asl; PEM A13845-6, PEM A14123 (tadpoles), Cuiva River source, -12.66825° 18.35282°, 1,407 m asl; PEM A12786, Cuando River source, trap Amphib. Reptile Conserv. 26 nN Fig. 5. Adult male Leptopelis sp. from Cuanavale River source. Photo by Luke Verburet. Fig. 6. Tadpole of Leptopelis sp. from Cuiva River source. Photo by Werner Conradie. Elevation (m) Map 5. Distribution of Leptopelis sp. in Angola. 4, -13.00164° 19.12960°, 1,361 m asl; PEM A12819- 21, Cuiva River source, -12.66856° 18.35307°, 1,433 m asl; PEM A14775, INBAC: WC-6852, Quembo River bridge camp, -13.52746° 19.28060°, 1,241 m asl; PEM A14767-—74, Quembo River right side tributary (Micongo River) past village, -13.51877° 19.28487°, 1,248 m asl. Additional material (3 specimens): P2- 277, wetland near old quarry east of Quemba, -12.16960° 18.22965°, 1,353 m asl; SAIAB 209098 (2 specimens), August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. Quembo River source lake, -13.14025° 19.04822°, 1,365 m asl. Description: Small terrestrial tree frog; large protruding eyes; tympanum clearly visible; well- developed rounded inner metatarsal tubercle; reduced webbing; digit tips slightly dilated (slightly wider than finger), finger tips more dilated than toe tips. Dorsum tan-brown; dark brown interorbital bar often present: dark brown vertebral stripe from just behind head to vent; some individuals with extra paravertebral bands; dark brown facial mask extending past arm onto side of body; scattered white speckles; groin with scattered unpigmented skin, extending onto the limbs; throat darkly pigmented; ventrum immaculate. Males with weakly developed pectoral glands and darkened throats. Adult females (n = 2) varied from 36.6—-40.7 (38.7) mm (largest female: PEM A12786); adult males (n = 35) varied from 29.0—34.6 (31.1) mm (largest male: PEM A12883). Elongated brown tadpoles; 40.7—52.2 mm total length, with tail 3.2 times body length; strong tail muscle starting just behind eye; thin fin margin above and below; LTRF 2(1)/3(1); jaw sheaths heavily keratinized, anterior part free of elongated marginal papillae. Habitat and natural history notes: Males were found calling in trees and shrubs (0.5—3 m above the ground), which were often located far from water, in grasslands with sparse tree cover. As the rainy season progressed, calls were heard progressively closer to water bodies, until eventually being heard among vegetation in the wetlands. The call resembles a chuckle. The eel-like tadpoles were found in wetlands among dense aquatic vegetation. Comments: Unusual light pink/red eel-like tadpoles were initially found in wetlands at the source of two different river systems, the Cuito and Kwanza rivers, in February 2016. A small number of these tadpoles were raised to adults in captivity. Nearly seven months were required to reach metamorphosis, during which time they changed from light pink/red to a more brownish coloration. They seemed to be sensitive to light as they swam eratically when removed from a dark environment and exposed to bright light. During the start of the rainy season in October 2016, unusual calls were heard which could not Fig. 7. Adult male Breviceps ombelanonga from Cuanavale River source. Photo by Luke Verburgt. Amphib. Reptile Conserv. 27 be assigned to a known species. After triangulation, several individuals were found calling from trees or shrubs, which were often far from water. On closer inspection, these specimens were found to be morphologically identical to the ones raised in captivity. We subsequently found these frogs at all major river sources and along rivers. Barcode analysis (16S rRNA) recovered the unknown Leptopelis as similar to L. ocellatus (94% similarity; KY080253), but the latter is a forest species with well-developed discs on toes. Schmidt and Inger (1959) described Leptopelis parvus from the Democratic Republic of the Congo (DRC), which resembles our Leptopelis sp. because of its small size and the shared absence of a white stripe above the vent that stretches onto the legs. However, our specimens differ from L. parvus in dorsal coloration pattern (para- and vertebral stripe present versus absent in L. parvus), lack of discs (dilated toe tips, but no clear discs versus clear discs in L. parvus), and pectoral glands (present versus absent in L. parvus). Further phylogenetic and morphological work is needed to resolve the taxonomic status of this species. Brevicipitidae Breviceps ombelanonga Nielsen, Conradie, Ceriaco, Bauer, Heinicke, Stanley, and Blackburn, 2020 Angolan Rain Frog (Fig. 7; Map 6) Material (4 specimens): PEM A12787, Quembo River source lake, -13.13544° 19.04397°, 1,374 m asl; PEM A12800, Cuanavale River source lake, -13.08934° 18.89485°, 1,359 m asl; PEM A12770, Cuando River source, -13.00334° 19.13564°, 1,362 m asl; PEM A12537, Cuito River source lake, -12.68935° 18.36012°, 1,431 m asl. Additional material (1 specimen): SAIAB 204537, Quembo River source lake, -13.13583° 19.04528°, 1,370 m asl. Description: Medium sized Breviceps; stout body; snout extremely shortened; pupils horizontally elliptical; tympanum not visible; outer metacarpal tubercle flat and undivided: short limbs; webbing absent; well-developed inner metatarsal tubercle fused with the outer metatarsal tubercle, with no deep cleft present, elongated, and Elevation (m) Map 6. Distribution of Breviceps ombelanonga in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola protruding outwards. Coloration varied from red with scattered black blotches, dark brown with red spots and markings, and light brown with red spots and black blotches (Nielsen et al. 2020). The only female collected measured 30.1 mm (PEM A12770); adult males (n = 3) varied from 18.3—26.6 (26.2) mm (largest male: PEM A12787). Habitat and natural history notes: Males were only heard calling during the day, especially after heavy rains; calling did not continue into the evenings. Call sites were among leaf litter in dense miombo woodland. Comments: The taxonomic status of Angolan Breviceps was recently addressed, leading to the description of this material as a new species, B. ombelanonga (Nielsen et al. 2020). Broader sampling across Angola may detect the presence of other species (e.g., B. adspersus to the south and B. poweri to the east) and even additional undescribed species. For now, all historical records are mapped as B. ombelanonga, until their taxonomic status can be confirmed. Bufonidae Sclerophrys gutturalis (Power, 1827) Guttural Toad (Fig. 8; Map 7) Material (69 specimens): PEM A12484, INBAC: WC-4841—2. Cuanavale River source lake, -13.08537° 18.89098°, 1,360 m asl; PEM A12498, drive back from Cuchi to Menongue, -14.67986° 17.17512°, 1,404 m asl; PEM A12573-—5, Cuito River source lake, -12.68935° 18.36012°, 1,431 masl; PEMA12612, Calua River source, 6 km SE of Cuito River source, -12.73675° 18.39310°, 1,442 m asl; PEM A12625-6, HALO Cuito , -12.39584° 16.96067°, 1,697 m asl; PEM A12629, roadside ditch 10 km SW of Cuito town, -12.44815° 16.88118°, 1,742 m asl; PEM A12639, PEM A12643, Cuanavale River, -13.37406° 18.99269°, 1,304 m asl; PEM A12677-9, INBAC: WC-5232, Camp 3, Malova Village, Mipanha River, -14.09140° 16.41476°, 1,553 masl; PEM A12685, Huambo HALO training camp, -12.73726° 15.81828°, 1,665 m asl; PEM A12698, Cuando River bridge, -13.60757° 19.53257°, 1,277 m asl; PEM A12719—25, Fig. 8. Adult male Sclerophrys gutturalis from Lungwebungu River source. Photo by Werner Conradie. Amphib. Reptile Conserv. Lungwebungu River camp bridge crossing, -12.58346° 18.66598°, 1,304 m asl; PEM A12796-7, Cuanavale River source lake camp side, -13.09442° 18.89372°, 1,368 m asl; PEM A12907-8, INBAC: no number (x2), Quembo River source lake, -13.13624° 19.04591°, 1,366 m asl; PEM A13763, Lungwebungu River, trap 2, -12.58199° 18.66562°, 1,208 m asl; PEM A13781, Lungwebungu River, trap 3, -12.58056° 18.66419°, 1,302 m asl; PEM A13784, Cuquema River, downstream, -12.47021° 16.82334°, 1,644 m asl, PEM A13790, Dam/ Hydroplant on Cuquema River, -12.42556° 16.81856°, 1,640 m asl; PEM A14680, Menongue, -14.63015° 17.63465°, 1,373 m asl; PEM A14685, wetland near old quarry east of Quemba, -12.16960° 18.22965°, 1,353 m asl; PEM A14712—5, INBAC: WC-6975, Lungwebungu River camp, -12.58439° 18.66748°, 1,297 m asl; PEM A14724, Quembo River bridge, -13.52746° 19.28060°, 1,241 m asl; PEM A14739, Quembo River bridge camp, trap 1, -13.52801° 19.28147°, 1,236 masl; PEM A14740, Quembo River bridge camp, -13.52746° 19.28060°, 1,241 m asl; PEM A14744, Quembo River bridge camp, trap 4, -13.52658° 19.27810°, 1,248 masl; PEMA14818— 22, INBAC: WC-7004, Luio River camp floodplains, -13.19711° 20.22194°, 1,181 m asl; PEM A14824-6, INBAC: WC-7029, Camp at side tributary (Luandai River) of the Luanguinga River, -13.70885° 21.26234°, 1,116 m asl; PEM A14831, Luanguinga River waterfall, -13.71132° 21.24914°, 1,118 m asl; PEM A14851-5, INBAC: WC-7044, Lake Hundo, -14.97431° 21.62966°, 1,100 m asl; PEM A14889-90, INBAC: WC-7079, Luvu River camp, -13.71200° 21.83538°, 1,082 m asl. Description: Large robust toad; snout rounded; elevated parotoid glands. Dorsum dark brown with pairs of dark paravertebral patches; pairs of dark patches on snout and behind eyes that create the appearance of a pale cross on head; in PEM A12724 and A12720 a continuous dark interorbital bar is present; back of thighs often with red infusions (not always present in females or juveniles). In breeding males, the dorsum becomes yellow, with numerous small black-tipped asperites; throat darkened; enlarged arms; black nuptial pads present on palm and Elevation (m) 3S o Oo Map 7. Distribution of Sclerophrys gutturalis in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. first finger. Adult females (n = 22) varied from 54.4— 109.7 (78.3) mm (largest female: PEM A12575); adult males (n = 37) varied from 56.7—89.2 (74.6) mm (largest male: PEM A14890). Habitat and natural history notes: Found in miombo woodland. Males were heard calling from margins of the source lakes. Comments: Widespread species across most of south-central Africa (Channing and Rédel 2019). Most historical Angolan material has been referred to as Bufo regularis Reus, 1833. Since the description of Bufo regularis gutturalis Power, 1927 (subsequently elevated to full species), the Angolan material has only partly been reassigned (Ruas 1996), and the rest of the extant material needs to be re- assessed to verify the taxonomic status. Additionally, this species needs to be reassessed as other authors have demonstrated that cryptic species may be present in this taxon (Pickersgill 2007; Telford et al. 2019). Sclerophrys pusilla (Mertens, 1937) Flat-backed Toad (Fig. 9; Map 8) Material (41 specimens): PEM A12425—6, INBAC (no number), Cunde waterfall, -13.77364° 18.75514°, 1,287 masl; PEMA12434, south of Menongue en route to Cuebe River, -14.96288° 17.69090°, 1,319 masl; PEM A12446, Cuchi River gorge, -14.59000° 16.90758°, 1,375 m asl; PEM A12494, HALO Menongue, -14.66317° 17.66521°, 1,386 m asl; PEM A12499, INBAC (no number), drive back from Cuchi to Menongue, -14.67986° 17.17512°, 1,404 m asl; PEM A12623-4, HALO Cuito , -12.39584° 16.96067°, 1,697 m asl; PEM A12630-1, roadside ditch 10 km SW of Cuito, -12.44815° 16.88118°, 1,742 m asl; PEM A12636, Quembo River source camp, -13.52653° 19.28368°, 1,242 m asl; PEM A12640, Cuanavale River, -13.37406° 18.99269°, 1,297 m asl; PEM A12642, Cuanavale River, -13.29236° 18.96283°, 1,313 m asl; PEM A 12650—1, Kwanza River bridge, -11.99348° 17.66965°, 1,273 m asl; PEM A12652, Kuvango Hydro Plant Site, wetland to east, -14.38755° 16.30166°, 1,451 m asl; PEM A12658, old Kuvango Hydroplant site, -14.38775° 16.29365°, 1,440 m asl; PEM A12659-60, Campsite 2 near Cuvango Mission, -13.32887 16.41167, 1,537 m asl; PEM A12668, Campsite 1 below rapids, Fig. 9. Adult male Sclerophrys pusilla from Cuito town. Photo by Werner Conradie. Amphib. Reptile Conserv. west of Fundo village, -13.04483° 16.37520°, 1,565 m asl; PEM A12680, Camp 3, Malova Village, Mipanha River, -14.09140° 16.41476°, 1,553 masl; PEM A12686- 7, Huambo HALO training camp, -12.73726° 15.81828°, 1,665 m asl; PEM A12690, Cubango 2017 launch site, -12.61700° 16.22132°, 1,727 m asl; PEM A12839, 31 km W of Menongue, Cueli River, -14.70511° 17.38014°, 1,392 masl; PEM A13791, Dam/Hydroplant on Cuquema River, -12.42556° 16.81856°, 1,640 m asl; PEM A14681— 2, Menongue, -14.63015° 17.63465°, 1,373 m asl; PEM A14720-1, 14725, Quembo River bridge, -13.52746° 19.28060°, 1,241 m asl; PEM A14745—7, Quembo River bridge camp, -13.52746° 19.28060°, 1,241 m asl; PEM A17761—2, Quembo River right side tributary (Micongo River) past village, -13.51877° 19.28487°, 1,478 m asl. Description: Medium sized toad; sharp snout profile; small flattened parotoid glands. Light brown dorsal coloration, with irregular darker markings; no interorbital bar; thin light dorsal stripe may be present; no red markings on the back of the thighs. Males with darkened throats; dorsum with black tipped asperites. Adult females (n = 16) varied from 52.1—79.3 (65.9) mm (largest female: PEM A14745); adult males (n = 9) varied from 35.5—58.7 (50.3) mm (largest male: PEM A12839). Habitat and natural history notes: Found in miombo woodland, sympatric with S. gutturalis. Comments: Poynton et al. (2016) recently split West Africa S. maculata from central and southern African S. pusilla. All the historical Angolan material referred to under the former name, S. maculata, now represents S. pusilla. Sclerophrys poweri (Hewitt, 1935) Western Olive Toad (Fig. 10; Map 9) Material (8 specimens): PEM A14876-82, INBAC: WC-7076, wetland south of Lake Hundo, -15.01099° 21.63608°, 1,100 m asl. Description: Large robust toad; snout rounded; large elevated parotoid glands. Dorsum of females tan with distinct black-edged dark brown to deep red paired dorsal markings; in males it can be more olive-yellow; red infusions on the back of the thighs. Males with dark throat and dorsal surface spinose with -12 Elevation (m) “14 -16 Map 8. Distribution of Sclerophrys pusilla in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola Fig. 10. Adult female Sclerophrys poweri from Lake Hundo. Photo by Werner Conradie. small black tipped asperites. Adult females (n = 3) varied from 101.0-102.3 (101.8) mm (largest female: PEM A14876; maximum recorded size); adult males (n = 5) varied from 85.5—97.0 (93.3) mm (largest male: PEM A14877; maximum recorded size). Habitat and natural history notes: This species was heard calling during early evenings in a flooded wetland and from margins of large natural lakes. Comments: Only a few records exist for southern and eastern Angola (Marques et al. 2018; Baptista et al. 2019). These specimens represent the easternmost records in Angola, and only the second record for Moxico Province (Ruas 1996). Hemisotidae Hemisus guineensis Cope, 1865 Guinea Shovel-snouted Frog (Fig. 11; Map 10) Material (6 specimens): PEM A14955, Cuando River, CUD2018 AC Camp 27, -16.09006° 21.83947°, 1,038 m asl; PEM A14832-3, INBAC: WC-6948, Lake Hundo, -14.97431° 21.62966°, 1,100 m asl: PEM A13831, Cuando River, camp 18, -14.66105° 20.16858°, 1,124 m asl; PEM A12771, Cuando River Source, trap 3, -13.00334° 19.13564°, 1,360 m asl. Additional material (1 tadpole lot): SAIAB 209095 (7 Fig. 11. Adult female Hemisus guineensis from Cuando River source. Photo by Werner Conradie. Amphib. Reptile Conserv. Elevation (m) 3 oO oO Map 9. Distribution of Sclerophrys poweri in Angola. tadpoles), small wooden bridge across wetland on road between Cuanavale River source camp and Munhango, -12.30714° 18.62333°, 1,397 m asl. Description: Small to medium sized frog; hardened pointed snout; small eyes; tympanum hidden; smooth dorsum (except PEM A12771, in which the yellow spots are slightly elevated): transverse skin ridge between posterior corners of eye, extending behind eye to above the arm; reduced webbing; large inner metatarsal and outer metacarpal tubercles. Grey dorsum with yellow mottling or spots; yellow vertebral stripe present; ventrum granular, with small irregular spots. Males with dark throats. Adult females (n= 3) varied from 40.5—49.2 (43.4) mm (largest female: PEM A12771); adult males (n= 3) varied from 26.7—31.6 (29.8) mm (largest male: PEM A14955). Habitat and natural history notes: No calls were heard. Specimens were either caught in traps or by hand while they were active at night after heavy rains in November, near open grassland and pans. One female (PEM A13831) collected in November was gravid. Comments: Laurent (1972) assigned all Angolan material he examined to the subspecies H. guineensis microps, and this was followed by Ruas (1996). However, Channing (2001) and Marques et al. (2018) documented two species of Hemisus occuring in Angola, Hemisus guineensis in the Elevation (m) 3 oO Oo Map 10. Distribution of Hemisus guineensis in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. Fig. 12. Adult female Hyperolius benguellensis from Cubango River rapids near Fundo village. Photo by Werner Conradie. north and H. marmoratus in the south-central region. We follow Laurent (1972) and assign all Angolan material to Hemisus guineensis until an in-depth phylogenetic work is conducted to assess the taxonomic status of the available material. Hyperoliidae Hyperolius benguellensis (Bocage, 1893) Benguela Reed Frog (Fig. 12; Map 11) Material (8 specimens, | tadpole lot): PEM A12438-41, INBAC (no number x2), Cuchi River gorge, -14.59000° 16.90758°, 1,365 m asl; PEM A12661, PEM A14172, Campsite 2 near old Cuvango Mission, -13.33451° 16.41280°, 1,356 masl; PEM A12675—6, Camp 3, Malova Village, Mipanha River, -14.09140° 16.41476°, 1,553 m asl; PEM A12666, Cubango River campsite 1 below rapids, west of Fundo village, -13.04790° 16.37896°, 1,568 m asl. Additional specimens (1 tadpole lot): SAIAB 209058 (18 tadpoles), Cuvango power station, entrance to canal, -14.38650° 16.28767°, 1,457 m asl. Description: Small reed frog; sharp but truncated snout, with small to no anterior protrusion; pedal webbing formula: I(1), I i/e (1- 0.5), TT (1-0.5), TV (1-1), V (0.75). Dorsum green with white dorso-lateral stripes (mostly males) or uniformly green with scattered brown spots (mostly females); ventrum transparent. Males with yellow gular disc. Single adult female measured 24.7 mm (PEM A12438); adult males (n = 7) varied from 16.2—22.8 (19.7) mm (largest male: PEM A12440). Habitat and natural history notes: Found on margins of rivers and in wetlands. Restricted to the western side of the study area, associated with the Cubango River system. Comments: In the most recent revision of the Hyperolius nasutus-complex, 16 species have been recognized (Channing et al. 2013). At least four species are expected to occur in Angola (Channing et al. 2013; Marques et al. 2018; Baptista et al. 2019), namely H. benguellensis, H. nasutus, H. adspersus, and H. dartevellei. Two clear morphotypes exist based on the general snout shape: sharp (benguellensis group) and Amphib. Reptile Conserv. 31 Elevation (m) 12 14 16 18 20 Map 11. Distribution of Hyperolius benguellensis in Angola. rounded (nasutus group). Of the sharp snouted form, we distinguished between two morphotypes in southeastern Angola: the ‘shark’-like profile (H. benguellensis, this Species account) and the acutely pointed snout, with a distinct protruding tip (H. cf. inyangae, see species account below). These identifications were confirmed by comparing 16S rRNA barcodes (W. Conradie, unpub. data) to published sequences (Channing et al. 2012). The remaining material is assigned to the nasutus group (see species account below). However, it must be noted that these little green frogs have been the subject of rigorous taxonomic debate over the years due to their cryptic nature, molecular and morphological similarity, and substantial geographic overlaps (see Channing et al. 2012 for overview). The taxonomic status of Angolan species belonging to these groups needs to be assessed in a broad- scale phylogenetic study. Hyperolius cinereus Monard, 1937 Ashy Reed Frog (Fig. 13; Map 12) Material (12 specimens, 1 tadpole lot): PEM A12442— 4, INBAC (no numbers x 2), Cuchi River gorge, -14.59000° 16.90758°, 1,375 m asl; PEM A12664, PEM A12670, Cubango River, campsite 1 below rapids, west of Fundo village, -13.04790° 16.37806°, 1,565 m asl; PEM A13787-9, INBAC: WC-520, Dam/ Hydroplant on Cuquema River, -12.42556° 16.81856°, 1,640 m asl; PEM A14128 (10 tadpoles), old Kuvango Hydroplant Site, wetland to east, -14.38755° 16.30166°, 1,438 m asl; INBAC: WC-5169, Cubango River source site, -12.66256° 16.09324°, 1,771 m asl. Description: Medium sized reed frog. No sexual dichromatism observed; both sexes with lime green to olive dorsum; protruding yellow eyes; red inner thighs; ventrum yellow. Males with yellow gular disc. Adult females (n = 3) varied from 20.3—29.1 (25.0) mm (largest female PEM A13789); adult males (n = 9) varied from 19.1— 22.8 (21.6) mm (largest male: PEM A12443). Habitat and natural history notes: Only recorded from the western side of the study area, where it was associated August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola Fig. 13. Adult Hyperolius cinereus from Cubango River rapids near Fundo village. Photo by Werner Conradie. with the Cubango and Cuito rivers. Comments: Widely recorded from the interior highlands of Angola (Conradie et al. 2013). The map in Marques et al. (2018) incorrectly plotted records from southern Cuando Cubango Province. The southeasternmost Angolan record of this species 1s close to the town of Menongue (Conradie et al. 2013). Hyperolius cf. inyvangae Laurent, 1943 Nyanga Reed Frog (Fig. 14; Map 13) Material (30 specimens): PEM A12730-3, INBAC: WC-4839; INBAC (no number x3), Cuando River source, -13.00346° 19.12751°, 1,353 m asl; PEM A14793-— 803, INBAC: WC-7023, INBAC: WC-7025, Luio River camp floodplains, -13.19711° 20.22194°, 1,181 m asl; PEM A13741, Lungwebungu River campsite, -12.58319° 18.66570°, 1,284 m asl; PEM A14887-8, Luvu River camp, -13.71200° 21.83538°, 1,082 m asl: PEM A12858—9, Quembo River source lake, -13.13624° 19.04591°, 1,366 m asl; PEM A12500, PEM A12513, river crossing before Samboano Village, -12.30700° 18.62350°, 1,398 m asl; PEM A14892-3, wetland near old quarry east of Quemba, -12.16960° 18.22965°, 1,353 m asl. Additional material (1 specimen): P2- Fig. 14. Adult male Hyperolius cf. invangae from Cuando River source. Photo by Werner Conradie. Amphib. Reptile Conserv. — 2000 - 1500 — 1000 Elevation (m) Map 12. Distribution of Hyperolius cinereus in Angola. 274 (photograph and tissue sample), wetland near old quarry east of Quemba, -12.16960° 18.22965°, 1,353 m asl. Description: Small slender species of reed frog; very sharp snout with a small anterior protrusion, which extends well beyond the margin of the mouth; small black asperites on throat; pedal webbing formula: I(1), II w/e (1-0.75), IM (1-0.75), TV (0.75-0.75), V (0.5). Dorsum lime green with white dorsolateral stripes; ventrum transparent; toe tips and webbing yellow. Adult females (n= 9) varied from 15.0—19.6 (16.7) mm (largest female: PEM A12500); adult males (n = 21) varied from 12.8-17.9 (15.1) mm (largest male: PEM A14893). Habitat and natural history notes: Found sympatric with Hyperolius nasutus at the Lungwebungu, Cuando, and Quembo rivers, where they were distinguished by microhabitat preference. Hyperolius cf. invangae was found among vegetation associated with slow running water, while H. nasutus was found on margins of source lakes or flooded areas with larger bodies of open water. Comments: Bittencourt-Silva (2019) assigned a specimen from western Zambia to H. nasicus based on head shape and webbing, but mentioned that molecularly itis most like H. invangae. The new material documented Elevation (m) Map 13. Distribution of Hyperolius cf. invangae in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. here is molecularly identical (16S rRNA: 97.5—100%) to the published sequences of H. inyangae (Channing et al. 2013; Bittencourt-Silva 2019) and agrees with the description, especially regarding the sharp snout usually with a pointed protrusion, but differs in the pedal webbing condition (Channing et al. 2013). It is reported to have reduced webbing and is illustrated as such, while the specimens here have more extensive webbing. This is either an error or there is a degree of variation in the webbing condition. The presence of this species so far to the west is an unexpected result, as H. invangae is currently only known from the Eastern Highlands of Zimbabwe. These new records thus represent a range extension of over 1,500 km westward. Further phylogenetic work is underway to fully document the taxonomic status of this population and other species assigned to the H. nasutus or H. benguellensis complexes. Hyperolius nasutus Gunther, 1865 Large-nosed Reed Frog (Fig. 15; Map 14) Material (64 specimens, 1 tadpole lot): PEM A12599-601, Calua River source, 6 km SE of Cuito River source, -12.73675° 18.39310°, 1,445 m asl: PEM A12424, Cunde waterfall, -13.77390° 18.75520°, 1,285 m asl; PEM A14107 (tadpoles), PEM A12435, confluence of Cuito and Calua rivers, -13.12458° 18.20909°, 1,345 m asl; PEM A12461-3, INBAC (no number x 4), Cuanavale River source lake, -13.08537° 18.89098°, 1,360 m asl; PEM A12738—40, INBAC (no number), Cuando River source, -13.00346° 19.12751°, 1,353 m asl; PEM A12693, Cubango River source site, -12.66256° 16.09324°, 1,771 m asl; PEM A12445, Cuchi River gorge, -14.5900° 16.90758°, 1,365 m asl; PEM A12550-1, Cuito River source lake, -12.68935° 18.36012°, 1,431 m asl; PEM A12427, INBAC (no number), Cuiva River bridge on EN250, -11.98345° 17.72367°, 1,267 m asl; PEM A12490, Dala River, near Samanunga village, -12.93169° 18.81458°, 1,363 m asl; PEM A12665, Cubango River campsite 1 below Fig. 15. Adult male Hyperolius nasutus from Cuanavale River source. Photo by Werner Conradie. Amphib. Reptile Conserv. rapids, west of Fundo village, -13.04790° 16.37896°, 1,568 m asl; PEM A12805-6, PEM A13804—12, Lake Tchanssengwe, -12.41402° 18.64418°, 1,393 m_ asl; PEM A14827-8, INBAC (no number), Luanguinga River waterfall, -13.71132° 21.24914°, 1,118 m asl; PEM A13773-80, Lungwebungu River old oxbows, -12.58129° 18.67162°, 1,304 m asl; PEM A14750—52, Quembo River right side tributary (Micongo River) past village, -13.51877° 19.28487°, 1,248 m asl; PEM A12860-62, INBAC (no number), Quembo River source lake, -13.13624° 19.04591°, 1,366 m asl; PEM A14729-35, INBAC (no number x2), Quembo River, oxbow near small waterfall, -13.54257° 19.29551°, 1,233 m asl. Additional material (13 specimens, 8 tadpole lots): SATAB 209057 (1 specimen), Cuanavale River near confluence, -13.12478° 18.90017°, 1,346 m asl; SAIAB 204573 (2 specimens), Quembo River source lake, -13.13586° 19.04492°, 1,369 m asl; SAIAB 209053 (7 tadpoles), outlet River from Kuembo River source lake, -13.13689° 19.03144°, 1,392 m asl; SAIAB 209173 (3 tadpoles), Quembo River source, -13.13583° 19.04528°, 1,370 m asl; SAIAB 209051 (5 specimens), Quembo River source lake, -13.14025° 19.04822°, 1,365 m asl; SAIAB 209045 (6 tadpoles), stream outflow about 2.6 km downstream of source lake, -13.00317° 19.15153°, 1,331 m asl; SAIAB 209056 (3 specimens), SAIAB 209087 (1 specimen), small stream 3 km below Cuanavale River source camp, -13.12539° 18.89914°, 1,344 m asl; SAIAB 209049 (5 tadpoles), south west of Cambuta on main track, -13.44678° 19.96403°, 1,229 m asl; SAIAB 209055 (1 tadpole), river bridge at Cangamba, -13.69611° 19.87503°, 1,193 m asl; SAIAB 209050 (1 tadpole), Calua lagoon, -12.73600° 18.39394°, 1,448 m asl; SAIAB 209026 (1 tadpole), Cuanavale River lake outlet, -13.09414° 18.89612°, 1,357 m asl; SAIAB 209054 (4 tadpoles), Cuando River source bog, -13.00383° 19.12719°, 1,350 m asl. Description: Small reed frog; rounded snout. Dorsum green, with white dorso-lateral stripes and small scattered black spots; Elevation (m) ° i=) oO Map 14. Distribution of Hyperolius nasutus in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola ventrum transparent; pedal webbing formula: I(1), II i/e (1-0.25), TT (0.75-0.5), IV (0.75-0.5), V (0.5). Male throat is white. Adult females (n = 14) varied from 15.9— 20.2 (18.5) mm (largest female: PEM A13810); adult males (n = 21) varied from 13.7—20.3 (17.2) mm (largest male: INBAC no number). Habitat and natural history notes: Found on the margins of source lakes, larger rivers, and cut-off oxbows. Comments: Found at certain localities in sympatry with Hyperolius cf. invangae (see above), but can easily be distinguished based on the rounded snout and larger overall size. Hyperolius parallelus Gunther, 1858 Angolan Reed Frog (Fig. 16; Map 15) Material (80 specimens, 6 tadpole lots): PEM A12436- 7, INBAC (no number), Cuchi River gorge, -14.59000° 16.90758°, 1,365 m asl; PEM A12448—-54, Cuanavale River source lake, -13.08537° 18.89098°, 1,360 m asl: PEM A12488, Dala River, near Samanunga village, -12.93169° 18.81458°, 1,363 m asl; PEM A12525, Muhango village, -12.16310° 18.55430°, 1,430 m asl: PEM A12520, PEM A12538-42, INBAC (no number), Cuito River source lake, -12.68935° 18.36012°, 1,431 m asl; PEM A12586—-91, PEM A12827, Calua River source lake, 6 km SE of Cuito River source, -12.73675° 18.39310°, 1,445 masl; PEM A12683—-4, Huambo HALO training camp, -12.73726° 15.81828°, 1,665 m asl; PEM A12851-—56, INBAC: WC-4626, INBAC (no number x2), Quembo River source lake, -13.13624° 19.04591°, 1,366 masl; PEM A13737 (tadpoles), Comba River, -12.62442° 18.65159°, 1,299 m asl; PEM A13765—6, Lungwebungu River, old oxbows, -12.58129° 18.67162°, 1,304 m asl; PEM A13794—8, PEM A13799 (tadpoles), INBAC: WC-4555, Lake Tchanssengwe, -12.41402° 18.64418°, 1,393 m asl; PEM A14097 (tadpoles), Dala River, near Samanunga village, -12.93169° 18.18146°, 1,315 m asl; PEM A14105 (tadpoles), Confluence of Cuito and Calua rivers, -13.12458° 18.89989°, 1,345 m asl; PEM A14113 (tadpoles), Cuiva River, -11.98346° 17.72841°, 1,264 m asl; PEM A14683—4, wetland near old quarry Fig. 16. Adult male Hyperolius parallelus from Cuanavale River source. Photo by Werner Conradie. Amphib. Reptile Conserv. 34 east of Quemba, -12.16960° 18.22965°, 1,353 m asl; PEM A14698-9, wetland west of Lungwebungu River camp, -12.55855° 18.6377°, 1,308 m asl; PEM A14726, Quembo River, oxbow near small waterfall, -13.54257° 19.29551°, 1,233 m asl; PEM A14783—92, INBAC: WC-7012, INBAC: WC-7008, INBAC: WC-7014, Luio River camp floodplains, -13.19711° 20.22194°, 1,181 m asl; PEM A14834—-43, INBAC: WC-6928, INBAC: WC-6928, Lake Hundo, -14.97431° 21.62966°, 1,100 m asl; PEM A14883—5, PEM A14886 (tadpoles), INBAC: WC-7083, Luvu River camp, -13.712° 21.83538°, 1,082 m asl; INBAC (no number), Cuiva River bridge on EN250, -11.98345° 17.72367°, 1,267 m asl. Additional material (1 specimen, 14 tadpole lots): SATAB 204515 (9 tadpoles), below the outlet of the Cuanavale River source lake, -13.09364° 18.89597°, 1,357 m asl; SAIAB 209030 (10 tadpoles), small bridge on road to Cuanavale River source, -12.30714° 18.62333°, 1,399 m asl; SAIAB 204563 (5 tadpoles), SAIAB 204566 (1 tadpole), Quembo River source lake, -13.13611° 19.04500°, 1,367 m asl; SAIAB 209025 (1 tadpole), Quembo River source lake outlet , -13.14025° 19.04822°, 1,365 m asl; SAIAB 209027 (1 tadpole), Cunde waterfall, -13.77364° 18.75514°, 1,287 m asl; SAIAB 209065 (8 tadpoles), swamp near Cuanavale River source, -13.10750° 18.86089°, 1,386 m asl; SAIAB 209085 (1 specimen), Cuanavale River source lake , -13.08997° 18.89561°, 1,358 m asl; SAIAB 204471 (16 tadpoles), frog pan 30 km below Cuando River source camp, -13.06831° 19.34369°, 1,297 m asl; SAIAB 209024 (1 tadpole), Samununga village, -12.93228° 18.81672°, 1,364 m asl; SAIAB 209029 (3 tadpoles), south west of Cambuta on main track, -13.44678° 19.96403°, 1,229 m asl; SAIAB 209028 (4 tadpoles), Cueve River source, peat bog source, -12.66949° 18.35203°, 1,420 m asl; SAIAB 209034 (5 tadpoles), Calua River lagoon, -12.73600° 18.39394°, 1,448 m asl; SAIAB 208959 (7 tadpoles), Cuanavale River source lake outlet, -13.09414° 18.89612°, 1,357 m asl; SAIAB 209031 (6 tadpoles), Cuanavale River bog above lake, -13.08575° 18.89215°, -14 Elevation (m) -16 -18 Map 15. Distribution of Hyperolius parallelus in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. 1,356 masl; SAIAB 209035 (7 tadpoles), Cuvango River - Power station camp, -14.38720° 16.28760°, 1,456 m asl. Description: Large reed frog. Dorsal coloration varied from finely vermiculated to boldly patterned: base color mostly orange to brick red with irregular black-edged white-cream markings that were often fused to form irregular stripes, the centers of these markings exhibited a small red or yellow spot or formed a thin line; feet and webbing red; ventrum white. Adult females (n = 24) varied from 26.8—37.2 (32.2) mm (largest female: PEM A14789); adult males (n = 52) varied from 22.8— 34.7 (29.2) mm (largest male: PEM A12542). Habitat and natural history notes: All specimens were found around larger water bodies. Males started calling in the early evening, high up in trees around the water bodies, and slowly moved to the water edge during the evening. Conradie et al. (2021) recorded predation of this species by the following species of snakes: Crotaphopeltis hotamboeia, Philothamnus ornatus, Philothamnus semivariegatus, and Kladirostratus acutus. Comments: This species was by far the most common frog from all major waterbodies surveyed. As in the larger viridiflavus group, many color variations have been described in the parallelus subgroup (see Channing 2022). The material from this study conforms to the color pattern recorded for Hyperolius angolensis Steindachner, 1867, which is now a synonym of H. parallelus. Hyperolius raymondi Conradie, Branch, and Tolley, 2013 Raymond’s Reed Frog (Fig. 17; Map 16) Material (27 specimens, 3 tadpole lots): PEM A12464—7, INBAC (no number x 2), Cuanavale River source lake, -13.08537° 18.89098°, 1,360 m asl; PEM A12514-8, river crossing before Samboano village, -12.30700° 18.62350°, 1,398 m asl; PEM A12521-2, PEM A14124 (tadpoles), Cuiva River source, -12.66825° 18.35282°, 1,407 m asl; PEM A12553-8, INBAC (no number x2), Cuito River source lake, -12.68935° 18.36012°, 1,431 m asl; PEM A12602-3, PEM A14120 (tadpoles), Calua River source lake, 6 km SE of Cuito River source, -12.73675° 18.39310°, 1,445 m asl; PEM Fig. 17. Adult male (above) and female (below) Hyperolius raymondi from river crossing before Samboano village. Photo by Luke Verburgt. Amphib. Reptile Conserv. A12741, Cuando River source, -13.00346° 19.12751°, 1,353 m asl; PEM A12816, Cuiva River source, -12.66856° 18.35307°, 1,433 m asl; PEM A13742-3, Lungwebungu River campsite, -12.58319° 18.66573°, 1,284 m asl; PEM A14127 (tadpoles), confluence of Cuito and Calua rivers, -12.74878° 18.35433°, 1,393 m asl. Additional material (1 specimen, 1 tadpole lot): P2-275 (photograph and tissue sample), wetland near old quarry east of Quemba, -12.16960° 18.22965°, 1,353 m asl; SAIAB 209041 (2 tadpoles), Quembo River source lake, -13.13586° 19.04492°, 1,372 m asl. Description: Medium sized reed frog. Most males possess two paravertebral and two dorsolateral black stripes on lime green to olive background; three males (PEM A12515, A13742-3) did not exhibit any dorsal stripes, and in one male (PEM A12816) the stripes faded mid-dorsally and eventually disappeared. All females devoid of dorsal markings. Adult females (n = 3) varied from 21.2—23.7 (22.6) mm (largest female: PEM A12555); adult males (n = 19) varied from 16.5—21.4 (18.8) mm (largest male: PEM A12816). Habitat and natural history notes: Found among inundated tall grass next to slow flowing water. Comments: This recently described species is currently regarded as endemic to Angola, but is expected to occur in adjacent DRC (Conradie et al. 2013). Initially, it was only known from the rivers draining northwards into the Congo basin (Conradie et al. 2013), and it was therefore thought to be associated with the Congo fauna. The discovery of this species in southeastern Angola corroborates Cei’s (1977) findings that the amphibian fauna located within the Cubango River system and has affinities with Congo fauna, due to the apparent lack of natural barriers between these regions. This is the southernmost record for Angola for this species, and the first from the Okavango and Zambezi River catchments. Hyperolius quinquevittatus Bocage, 1866 Five-striped Reed Frog (Fig. 18; Map 17) Material (1 specimen): P2-273 (photograph and tissue sample only), wetland near old quarry east of Quemba, -12.16960° 18.22965°, 1,353 m asl. Description: Elevation (m) 3 So oO Map 16. Distribution of Hyperolius raymondi in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola Fig. 18. Adult male Hyperolius quinquevittatus from wetland east of Quemba town. Photo by Pedro Vaz Pinto. Medium sized reed frog; pointed snout; long slender limbs. Dorsum dark brown, with a single vertebral and two dorsolateral light green stripes. Habitat and natural history notes: Found in a flooded grassland, surrounded by miombo woodland. Founded syntopically with Kassinula wittei, Hyperolius raymondi, and H. cf. invangae. Comments: This record represents an eastward range extension in Angola. It has been recorded from northwestern Zambia (Poynton and Broadley 1987, 1991; Channing 2001; Bittencourt 2019), and like Kassinula wittei and Kassina kuvangensis, this species 1s thus expected to occur to the north and east of the defined study area. Hyperolius aff. bocagei Steindachner, 1867 Bocage’s Reed Frog (Figs. 19-21; Map 18) Material (74 specimens, 10 tadpole lots): PEM A12423, Cunde waterfall, -13.77390° 18.75520°, 1,287 m asl; PEM A12455-60, PEM A14085-6 (tadpoles) Cuanavale River source lake, -13.08537° 18.89098°, 1,340 m asl; PEM A12489, PEM A14098 (tadpoles), INBAC (no number), Dala River, near Samanunga village, -12.93169° 18.81458°, 1,363 m asl; PEM A12508-12, river crossing before Samboano village, -12.30700° 18.62350°, 1,397 m asl; PEM A12543-52, INBAC Elevation (m) S ° Oo Map 18. Distribution of Hyperolius aff. bocagei in Angola. Amphib. Reptile Conserv. Elevation (m) I ra Oo So Fig. 19. Adult male Hyperolius aff. bocagei (green form) from Cuito River source. Photo by Werner Conradie. Fig. 20. Adult male Hyperolius aff. bocagei (red form) from Cuanavale River source. Photo by Werner Conradie. Fig. 21. Adult male Hyperolius aff. bocagei (brown form) from Cuito River source. Photo by Werner Conradie. August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. (no number x2), Cuito River source lake, -12.68935° 18.36012°, 1,431 m asl; PEM A12592-8, Calua River source lake, 6 km SE of Cuito River source, -12.73675° 18.39310°, 1,446 m asl; PEM A12619, Stop 1: road to Cuito River source, -12.25050° 18.63730°, 1,556 m asl; PEM A12674, Camp 3, Malova Village, Mipanha River, -14.09140° 16.41476°, 1,553 m asl; PEM A12734-7, INBAC: WC-4833, INBAC (no number x2), Cuando River source, -13.00346° 19.12751°, 1,553 m asl; PEM A12788-9, INBAC: WC-4614, Quembo River, source trap 2, -13.13544° 19.04397°, 1,374 m asl; PEM A14112 (tadpoles), Cuiva River, -11.98346° 17.72841°, 1,264 m asl; PEM A12815, PEM A14125 (tadpoles), Cuiva River source, -12.66856° 18.35307°, 1,407 m asl; PEM A12857, INBAC: WC-4616; INBAC (no number x5) Quembo River source lake, -13.13624° 19.04591°, 1,411 m asl; PEM A13767—72, INBAC: WC-6979 Lungwebungu River, old oxbows, -12.58129° 18.67162°, 1,304 m asl; PEM A13800—2, PEM A13803 (tadpoles), Lake Tchanssengwe, -12.41402° 18.64417°, 1,415 masl; PEM A14090 (tadpoles), 4 km upstream from Cuanavale River source, -13.05084° 18.89726°, 1,394 m asl; PEM A14103 (tadpoles), 4 km downstream from Cuanavale River source camp, -13.11585° 18.90246°, 1,354 m asl; PEM A14106 (tadpoles), confluence of Cuito and Calua rivers, -13.12458° 18.89989°, 1,345 m asl; PEM A14686, detour route just across bridge over Cuiva River, -12.13942° 18.39393°, 1,385 m asl; PEM A14693 (tadpoles), Lungwebungu River camp, first oxbow on right side, -12.58117° 18.67106°, 1,294 m asl; PEM A14717, en route from Samanunga village to Cuanavale River source, -12.94331° 18.81118°, 1,407 m asl; PEM A14727-8, Quembo River, oxbow near small waterfall, -13.54257° 19.29551°, 1,233 m asl; PEM A14748-9, PEM A14763—65, PEM A14778, Quembo River right side tributary (Micongo River) past village, -13.51877° 19.28487°, 1,248 m asl; PEM A14757, left side tributary (Condinde River) at Cuando River bridge, -13.60076° 19.52675° 1,219 m asl. Additional material (16 tadpole lots): SAIAB 204553 (7 tadpoles), Cuanavale River source lake, -13.08997° 18.89389°, 1,358 m asl: SAIAB 209059 (15 tadpoles), Cuanavale River near confluence, -13.12478° 18.90017°, 1,347 m asl; SAIAB 204509 (29 tadpoles), outlet from Kuembo River source lake, -13.13689° 19.03144°, 1,375 m asl; SAITAB 204564 (25 tadpoles), SAIAB 209074 (1 tadpole), Quembo River source lake, -13.13611° 19.04500°, 1,366 m asl; SAIAB 209066 (9 tadpoles), Quembo River source lake, -13.14025° 19.04822°, 1,365 m asl; SAIAB 209062 (2 tadpoles), swamp near Cuanavale River source lake camp, -13.10750° 18.86089°, 1,386 m asl; SAIAB 204465 (19 tadpoles), Cuando River source pool, -13.00383° 19.12719°, 1,350 m asl; SAIAB 209060 (2 tadpoles), stream outflow about 2.6 km downstream of source lake, -13.00317° 19.15153°, 1,331 m asl; SAIAB 204472 (13 tadpoles), Cuando River, -13.09142° 19.35850°, 1,283 m asl; SAIAB 209067 (8 tadpoles), on Cuanavale River Amphib. Reptile Conserv. road, -13.14190° 19.44503°, 1,301 masl; SATAB 209070 (10 tadpoles), pan near Cuanavale River camp, seepage, -13.10750° 18.86089°, 1,387 m asl; SAIAB 204569 (13 tadpoles), Samununga village, -12.93228° 18.81672°, 1,365 m asl; SAIAB 209063 (6 tadpoles), small stream 3 km below Cuanavale River source camp, -13.12539° 18.89914°, 1,344 m asl; SATAB 209069 (11 tadpoles), pool in wetland on road edge, west of Munhango, -12.17806° 18.24306°, 1,370 m asl; SAIAB 209075 (18 tadpoles), Cuando River source bog, -13.00383° 19.12719°, 1,350 masl. Description: Medium sized reed frog; rounded snout; reduced webbing; ventrum granular. Three different color forms were observed, 1.e., plain light to dark green, brown, and bright red; thin dorsolateral white stripe present; small white specks on dorsal side of legs; toes and webbing red. Tadpoles with ventrum spotted and ventral blotches remain present in juveniles but fade in adults. Adult females (n = 11) varied from 16.7-21.9 (19.6) mm (largest female: PEM A12549),; adult males (n = 52) varied from 14.8—21.1 (18.7) mm (largest male: PEM A12592). Habitat and natural history notes: Found at the margins of large rivers and source lakes within dense emergent wetland vegetation. Sympatric with other species of Hyperolius, mostly H. nasutus and H. parallelus. Comments: Initially we assigned these specimens to H. bocagei (NGOWP 2017), however it 1s generally regarded as a larger species that is associated with the viridiflavus group (see Schietz 1999). This new material differs in size (smaller) and webbing (less webbing) from H. bocagei (Schietz 1999). Phylogenetic work is currently underway to determine its taxonomic status. Kassina kuvangensis (Monard, 1937) Kuvangu Kassina (Fig. 22; Map 19) Material (9 specimens, 5 tadpole lots): PEM A12496- 7, unnamed side-triburaty source of Cuanavale River, -13.07518° 18.88481°, 1,374 m asl; PEM A12775-6, Quembo River, trap 4, -13.13586° 19.04709°, 1,369 m asl; PEM A12781, Quembo River trap 1, -13.13592° 19.04417°, 1,369 m asl; PEM A12825, Quembo River, trap 3, -13.13073° 19.03725°, 1,445 m asl; PEM A14116 (tadpoles), river crossing before Sombanana village, -12.30710° 18.62350°, 1,407 m asl; PEM A12828, PEM A14119 (tadpoles), Calua River source 6 km SE of Cuito River source, -12.73675° 18.39310°, 1,446 m asl; PEM A14089 (tadpoles), 4 km upstream from Cuanavale River source, -13.05084° 18.89726°, 1,394 m asl; PEM A14096, Dala River, near Samanga village, -12.93169° 18.81458°, 1,363 m asl; PEM A14101 (tadpoles), 4 km downstream from Cuanavale River source camp, -13.11585° 18.90246°, 1,354 m asl; PEM A14104 (tadpoles), confluence of Cuito and Calua rivers, -13.12458° 18.89989°, 1,345 m asl; PEM A14804, Luio River camp floodplains, -13.19711° 20.22194°, 1,181 m asl. Additional material (4 tadpole lots): SAIAB 209127 (1 tadpole), SAIAB 209107 (2 tadpoles), swamp August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola Fig. 22. Adult female Kassina kuvangensis from Cuanavale River source. Photo by Werner Conradie. near Cuanavale River lake camp, -13.10750° 18.86089°, 1,386 masl; SATAB 209108 (22 tadpoles), pool in wetland on road edge west of Munhango, -12.17806° 18.24306°, 1,381 m asl; SAIAB 209089 (4 tadpoles), Calua lagoon, -12.73599° 18.39394°, 1,448 m asl. Description: Large sized Kassina; tympanum visible; elliptical vertical pupils; large inner metatarsal tubercle; subarticular tubercle well-developed. Dorsum very dark olive-brown, with scattered large darker brown yellow-edged irregular shaped spots; ventrum yellow, but in some individuals has black-edged white/yellow spots. In females, the cloaca sides are swollen. In males, the gular flap is dark and the glands behind eye/head are enlarged. Adult females (n = 2) varied from 51.9-53.3 (52.6) mm (largest female: PEM A14096); adult males (n = 7) varied from 38.4—52.9 (43.7) mm (largest male: PEM A12781). Tadpoles can get very large (up to 135 mm total length); LTRF 1/2(1), with strong jaw sheaths. Habitat and natural history notes: In the evenings, males call from the margins of source lakes and flooded areas while clinging to vegetation, and quickly submerge themselves when disturbed. Comments: These new records and other records from Uige Province, which are the northwesternmost records (Ernst et al. 2020), represent the first adult K. Auvangensis Fig. 23. Adult female Kassina senegalensis from Culua River source. Photo by Werner Conradie. Amphib. Reptile Conserv. | " | I Elevation (m) ° Oo oO eh “14 -16 -18 Map 19. Distribution of Kassina kuvangensis in Angola. for Angola since the original species description (Monard 1937). The new localities recorded here lie between the Species type locality in Angola (Kuvango, Monard 1937) and western Zambia (Poynton and Broadley 1987, 1991; Channing 2001). Kassina senegalensis (Dumeéril and Bibron, 1841) Bubbling Kassina (Fig. 23; Map 20) Material (56 specimens, 3 tadpole lots): PEM A12604—7, Calua River source, 6 km SE of Cuito River source, -12.73675° 18.39310°, 1,445 m asl; PEM A12621, Cutata River, -12.56916° 16.49334°, 1,647 m asl; PEM A12699—700, Lungwebungu River camp bridge crossing, -12.58347° 18.66598°, 1,304 m asl; PEM A12785, Cuando River source trap 4, -13.00164° 19.12960°, 1,374 m asl; PEM A12790, Quembo River trap 2, -13.13544° 19.04397°, 1,374 m asl; PEM A12829-30, Culua River source, -12.73723° 18.39340°, 1,444 m asl; PEM A12863-9, INBAC (no number x2), Quembo River source lake, -13.13624° 19.04591°, 1,366 m asl; PEM A13744 (tadpoles), Lungwebungu River campsite, -12.58319° 18.66573°, 1,284 m asl; PEM A13757-62, INBAC: WC-6261, Lungwebungu River, trap 2, -12.58199° 18.66562°, 1,208 m asl; PEM A14094 Elevation (m) S oO =) Map 20. Distribution of Kassina senegalensis in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. (tadpoles), series of pans south of Tempué, -13.55719° 18.85519°, 1,315 m asl; PEM A14109 (tadpoles), Cutata River, -12.56916° 16.49334°, 1,647 m asl; PEM A14687, PEM A14700, INBAC: WC-6759, Lungwebungu River camp, -12.58439° 18.66748°, 1,297 m asl; PEM A14718, PEM A14722, Quembo River bridge camp, -13.52746° 19.28060°, 1,241 m asl; PEM A14753-4, PEM A14766, Quembo River right side tributary (Micongo River) past village, -13.51877° 19.28487°, 1,248 m asl; PEM A14805—9, INBAC: WC-7017, Luio River camp floodplains, -13.19711° 20.22194°, 1,181 m asl; PEM A14844—-50, PEM A14867—72, INBAC: WC-6916, INBAC: WC-6917, INBAC: WC-6953, Lake Hundo, -14.97431° 21.62966°, 1,100 m asl. Additional material (1 tadpole lot): SAIAB 209106 (28 tadpoles), small wooden bridge across wetland on road between Cuanavale River source camp and Munhango, -12.30714° 18.62333°, 1,399 m asl. Description: Medium sized Kassina; elliptical vertical pupils; tympanum visible. Dorsum yellow-mustard, with irregular shaped dark brown blotches, sometimes fused to form a vertebral stripe; ventrum white. In females, the cloaca is swollen, with papillae pointed downward. Male throats are black. Adult females (n = 16) varied from 27.9-41.1 (37.6) mm (largest female: INBAC: WC-6759); adult males (n = 34) varied from 35.2-43.6 (39.2) mm (largest male: PEM A14809). Habitat and natural history notes: This species was found in sympatry with K. kuvangensis in the study area, although not syntopically. Kassina senegalensis was found in more open habitat or sparse vegetation, calling from the edges of waterbodies, while K. kuvangensis was only heard calling from well- vegetated floodplains and source lakes. Comments: Schietz (1999) divided material of K. senegalensis into different forms, based on the dorsal markings. As pointed out by Poynton and Broadley (1987), there is considerable overlap in these color forms between and within populations. Due to the large distributional range of this species from Senegal to South Africa (Channing and Rodel 2019), spanning many different habitats and f Elevation (m) 3 oO oOo Map 21. Distribution of Kassinula wittei in Angola. Amphib. Reptile Conserv. biomes, cryptic species are expected, and this species needs a broad-scale phylogeographic study. Kassinula wittei Laurent, 1940 De Witte’s Clicking Frog (Figs. 24—25; Map 21) Material (48 specimens): PEM A12468—-5, PEM A12792-3, INBAC (no number x2), Cuanavale River source lake, -13.08537° 18.89098°, 1,360 m asl; PEM A12807-—12, INBAC (no number x3), Cuito River source lake, -12.68727° 18.36067°, 1,423 m asl; PEM A12817— 8, Cuiva River source, -12.66856° 18.35307°, 1,433 m asl; PEM A12870—2, INBAC (no number x2), Quembo River source lake, -13.13624° 19.04591°, 1,366 m asl; PEM A14271-3, INBAC: WC-6743, Lungwebungu River camp, first oxbow on right side, -12.58117° 18.67106°, 1,294 m asl; PEM A14274—5, Cuanavale River source, -13.08537° 18.89098°, 1,357 m asl; PEM A14276—80, Quembo River right side tributary (Micongo River) past village, -13.51877° 19.28487°, 1,248 m asl; PEM A14281-4, INBAC: WC- 6958-60, Luio River camp floodplains, -13.20191° 20.22144°, 1,181 m asl; PEM A14270, PEM A14285, wetland near old quarry east of Quemba, -12.16960° 18.22965°, 1,353 m asl. Additional material (4 specimens): P2-278, P2-279, Fig. 24. Adult male Kassinula wittei from Cuanavale River source. Photo by Werner Conradie. Fig. 25. Adult male Kassinula cf. wittei from wetland east of Quemba town. Photo by Chad Keates. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola P2-280, P2-281, wetland near old quarry east of Quemba, -12.16960° 18.22965°, 1,353 m asl. Description: Minute hyperoliid species. See Conradie et al. (2020b) for a detailed description of this material. The only female measured 14.7 mm (PEM A14284); adult males varied from 12.3—22.0 (14.2) mm (largest male: PEM A14270). Habitat and natural history notes: Found in flooded grassland and peat wetlands. Comments: The occurrence of this species in northeastern Angola was expected from a biogeographical point of view, but these are the first records of the species for the country. These new records extend the known range of this species from northwestern Zambia and southern DRC to central Angola by more than 400 km (see Conradie et al. 2020b). The taxonomic status of this genus has recently been validated phylogenetically and the species is considered to be closely related to Afrixalus, although its exact systematic placement remains unclear (see Conradie et al. 2020b; Ne€as et al. 2022). Two specimens (PEM A14270, PEM A14285) differ in the dorsal coloration pattern (vertebral band broken and no darker stipples in lighter bands versus continuous dark vertebral band and lighter bands, with fine dark stipples), maximum size (22 mm versus 16.7 mm), and potential differences in habitat preference from other K. wittei collected. These two specimens are similar in appearance to a specimen collected from Congolo River, Luando Strict Nature Reserve (FHK091) (see Conradie et al. 2020b). While Ne€as et al. (2022) pointed out the relatively deep divergence between the specimen from Congolo River and the eastern Angolan and topotypic DRC samples, this difference was not considered in Conradie et al. (2020b). Preliminary unpublished genetic results (W. Conradie, unpub. data) show that the two specimens listed here agree genetically with the western Angolan sample (Congolo River) and may represent an undescribed cryptic species. These specimens occur syntopically with typical K. wittei east of Quemba, although collected from different habitats and different times of the year. Fig. 26. Adult female Phrynobatrachus mababiensis from Cutatu River. Photo by Werner Conradie. Amphib. Reptile Conserv. Phrynobatrachidae Phrynobatrachus mababiensis FitzSimons, 1932 complex Dwarf Puddle Frog (Fig. 26; Map 22) Material (14 specimens, 1 tadpole lot): PEM A12429, series of pans south of Tempué, -13.55719° 18.85519°, 1,315 masl; PEM A12608, Calua River source, 6 km SE of Cuito River source, -12.73675° 18.39310°, 1,445 m asl; PEM A12622, Cutata River, -12.56916° 16.49334°, 1,647 m asl; PEM A12662, Campsite 2 near old Cuvango Mission, -13.33451° 16.41280°, 1,542 m asl; PEM A12688, Cubango 2017 launch site, -12.61700° 16.22133°, 1,727 m asl; PEM A12777, Quembo River source, trap 4, -13.13586° 19.04709°, 1,373 m_ asl; PEM A12840-1, west of Cuito town on Aludungo rd, -12.28700° 16.81716°, 1,739 m asl; PEM A12889, Quembo River source lake, -13.13624° 19.04591°, 1,366 m asl; PEM A13739, Lungwebungu River, new campsite, -12.58445° 18.66538°, 1,308 m asl; PEM A14743, Quembo River bridge camp, trap 2, -13.52816° 19.28067°, 1,240 m asl; PEM A14779, Quembo River bridge camp, trap 1, -13.52801° 19.28147°, 1,236 m asl; PEM A14873, wetland south of Lake Hundo, -15.01099° 21.63608°, 1,100 m asl; PEM A14114 (tadpoles), river before Sombanana village, -12.30710° 18.62350°, 1,407 m asl; INBAC: WC-5221, Campsite 2 near old Cuvango Mission, -13.33451° 16.41280°, 1,542 m asl. Additional material (3 tadpole lots): SAIAB 209079 (14 tadpoles), road to Cuanavale River camp, -12.54990° 18.67444°, 1,333 m asl; SAIAB 209105 (10 tadpoles), swamp near Cuanavale River source lake camp, 1,386 m asl; SAIAB 209111 (9 tadpoles), small wooden bridge across wetland on road between Cuanavale River source camp and Munhango, -12.30714° 18.62333°, 1,399 m asl. Description: Small Phrynobatrachus; scattered small warts present on the back; up to three and a half phalanges of longest toe free of webbing. Dorsum light brown, with scattered darker brown markings; lower lip barred in black and white; ventrum varied from immaculate to mottled Elevation (m) S oOo oO Map 22. Distribution of Phrynobatrachus mababiensis complex in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. with black. Males with black throats. Adult females (n = 3) varied from 12.8—18.0 (16.2) mm (largest female: PEM A14743); adult males (n = 3) varied from 12.6— 17.2 (14.7) mm (largest male: PEM A14779). Habitat and natural history notes: Found on edges of rivers and source lakes among marginal vegetation. Comments: Numerous cryptic species have been documented within P. mababiensis as currently recognized (Zimkus and Schick 2010; Zimkus et al. 2010), and the new material recorded here may represent more than one species (N. Baptista et al., unpub. data). Until further phylogenetic studies are conducted, this material is all included in the P. mababiensis complex. Phrynobatrachus natalensis (Smith, 1849) Snoring Puddle Frog (Fig. 27; Map 23) Material (20 specimens): PEM A12637-8, Cuanavale River, -13.37406° 18.99269°, 1,297 m asl; PEM A12711, Lungwebungu River camp bridge crossing, -12.58347° 18.66598°, 1,304 m asl; PEM A12890-3, INBAC: WC- 4599, INBAC (no number) , Quembo River source lake, -13.13624° 19.04591°, 1,366 m asl; PEM A13738, Comba River, -12.62442° 18.65159°, 1,299 m asl: PEM A13745—8, Lungwebungu River old campsite, -12.58319° 18.66573°, 1,284 m asl; PEM A14690, PEM A14705-6, INBAC: WC-6740, Lungwebungu River camp, -12.58439° 18.66748°, 1,297 m asl; PEM A14707, wetland west of Lungwebungu River camp, -12.55855° 18.63770°, 1,308 m asl; PEM A14810—-1, Luio River camp floodplains, -13.19711° 20.22194°, 1,181 m asl. Additional material (1 specimen): SAIAB 209104 (1 specimen), swamp near Cuanavale River source lake camp, -13.10750° 18.86089°, 1,386 m asl. Description: Medium sized Phrynobatrachus; dorsum with scattered elevated tubercles; reduced webbing; heel spine present; small outer metatarsal tubercle; large inner metatarsal tubercle; ridge running along outer toe; small tarsal ridge: mid-tarsal tubercle present; well-developed elevated subarticular tubercle. Dorsum coloration varies from grey to brown and even green; ventrum white. Female throats Fig. 27. Adult male Phrynobatrachus natalensis from Luio River. Photo by Chad Keates. Amphib. Reptile Conserv. are speckled, while throats of males are uniformly grey to black. Adult females (n = 15) varied from 23.6—32.6 (28.1) mm (largest female: PEM A13745); adult males (n = 5) varied from 26.3—30.4 (28.2) mm (largest male: PEM A14705). Habitat and natural history notes: Found in flooded grasslands associated with miombo woodland. Comments: This species is widespread in Angola (Marques et al. 2018). Although there were no records for southeastern Angola prior to Conradie et al. (2016), they are widespread east of the Zambian border (Poynton and Broadley 1985b, 1991; Channing, 2001). This is another complex within Phrynobatrachus containing several cryptic species, and thus deserving of further investigation (Zimkus et al. 2010; Bittencourt- Silva 2019). Pipidae Xenopus petersii Bocage, 1895 Peters’ Clawed Frog (Fig. 28; Map 24) Material (40 specimens): PEM A11617—-9, PEM A12576—-84, INBAC (no number x 4), Cuito River source lake, -12.68935° 18.36012°, 1,431 m_ asl; PEM A12430-3, pans south of Tempue, -13.55719° 18.85519°, 1,315 m asl; PEM A12519, river crossing before Samboano village, -12.30700° 18.62350°, 1,397 m asl; PEM A12613—5, Calua River source, 6 km SE of Cuito River source, -12.73675° 18.39310°, 1,446 m asl; PEM A12634—5, roadside ditch 10 km SW of Cuito town, -12.44815° 16.88118°, 1,742 masl; PEM A12682, Camp 3, Malova Village, Mipanha River, -14.09140° 16.41476°, 1,553 m asl; PEM A12691—2, INBAC: WC-5175, Cubango River launch site, -12.61700° 16.22133°, 1,727 m asl; PEM A12694, Cubango River source site, -12.66256° 16.09324°, 1,771 m asl; PEM A12695, INBAC: WC-5173, New dam, Katchingo, -12.60587° 16.22003°, 1,373 m asl; PEM A12697, Chicala Choloanga roadside quarry, -12.63611° 16.04282°, 1,858 m asl; PEM A12779—-80, Quembo River, trap 4, -13.13586° 19.04709°, 1,373 m asl; Elevation (m) S (=) oO Map 23. Distribution of Phrynobatrachus natalensis in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola Fig. 28. Adult female Xenopus petersii from Cuito town. Photo by Werner Conradie. PEM A12909, Quembo River source lake, -13.13624° 19.04591°, 1,411 m asl; PEM A13756, Lungwebungu River old campsite, -12.58319° 18.66573°, 1,284 m asl: PEM A13817, river crossing before Samboano village, -12.20672° 18.06236°, 1,387 m asl; PEM A14776, Quembo River right side tributary (Micongo River) past village, -13.51877° 19.28487°, 1,248 m asl. Additional material (9 specimens, 1 tadpole lot): SAIAB 204517 (1 specimen), Lungwebungu River Bridge, -12.58397° 18.66536°, 1,295 m asl; SAIAB 204468 (1 tadpole), main road bridge over upper Kwanza east of cuito, -11.98433° 17.72197°, 1,267 m asl; SAIAB 204500 (1 specimen), small wooden bridge across wetland on road between Cuanavale source camp and Munhango, -12.30714° 18.62333°, 1,399 m asl; SAIAB 204502 (6 specimens), pool in wetland on road edge west of Munhango, -12.17806° 18.24306°, 1,370 m asl; SAIAB 209086 (1 specimen), Cuvango mission rapids camp, -13.32782° 16.41106°, 1,538 m asl. Description: Medium to large sized pipid; dorsum smooth; eyes on top of head; three clawed toes, no claw on prehallux: extensive webbing. Dorsum varies from light to dark brown; posterior half of ventrum and thighs with orange pigmentation. Adult females (n = 24) varied from 34.2— Elevation (m) Map 25. Distribution of Xenopus poweri in Angola. Amphib. Reptile Conserv. 42 Elevation (m) Map 24. Distribution of Xenopus petersii in Angola. 74.2 (52.4) mm (largest female: PEM A12682); adult males (n= 16) varied from 37.9-51.2 (45.5) mm (largest male: PEM A11619). Habitat and natural history notes: Aquatic species found in all major waterbodies surveyed. Comments: Furman et al. (2015) split X. petersii and X. poweri, restricting the former mostly to western Angola northward to Gabon. In Angola, this species seems to be absent from the south and east, where it is replaced by X. muelleri and/or _X. poweri. Xenopus poweri Hewitt, 1927 Powers’ Clawed Frog (Map 25) Material (13 specimens): PEM A14856-66, INBAC: WC-7040, INBAC: WC-7043, Lake Hundo, -14.97431° 21.62966°, 1,100 m asl. Description: Medium sized pipid; dorsum smooth; eyes on top of head; three clawed toes, no claw on prehallux; extensive webbing. Dorsum varies from light to dark brown, with scattered darker markings; ventrum grayish, with scattered darker blotches. Adult females (n = 6) varied from 55.6—65.9 (59.8) mm (largest female: PEM A14861); adult males (n = 6) varied from 43.1—51.2 (48.8) mm (largest male: PEM A14858). Habitat and natural history notes: After a heavy thunderstorm, numerous XY. poweri were found moving over land away from the waterbodies. Comments: Only recorded from lower elevations of eastern Angola, associated with open and dry savanna. Only a limited number of records exist of this species from eastern Angola (Conradie et al. 2016; Marques et al. 2018). Ptychadenidae Ptychadena bunoderma (Boulenger, 1907) Rough Ridged Frog (Fig. 29; Map 26) Material (2 specimens): PEM A12778, Quembo River source, trap 4, -13.13586° 19.04709°, 1,369 m asl; PEM A12476, Cuanavale River source lake, -13.08537° 18.89100°, 1,360 m asl. Description: Small stocky Ptychadena, dorsum warty, with no clear ridges as in other Ptychadena species; short stubby legs; very reduced August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. Fig. 29. Adult male Ptychadena bunoderma from Cuanavale River source lake. Photo by Werner Conradie. webbing, four phalanges of the 4" toe free of webbing. Dorsum dark brown to olive, with scattered white to brown elevated tubercles; lower lip gray with white bars; back of thighs with scattered light spots. The only adult male measured 25.6 mm (PEM A12778). Habitat and natural history notes: Specimens were found in trap arrays (see Conradie et al. 2021) set up next to flooded areas of source lakes. They were never heard calling, and either have a very cryptic call or were not vocal during the survey periods. Comments: In Angola, this species is only known from the east, based on historical records (Marques et al. 2018). No new collections were made until Ernst et al. (2020) recorded a series of specimens from northwestern Angola. Elsewhere, it is known from northwestern Zambia (Channing 2001; Channing and Rodel 2019). Our new material represents a new record for southeastern Angola and the Okavango River basin. Ptychadena grandisonae Laurent, 1954 Many-ridged Grass Frog (Fig. 30; Map 27) Material (3 specimens): PEM A12696, INBAC: WC- 5185, Chicala Choloanga roadside quarry, -12.63611° 16.04282°, 1,858 m asl; PEM A12689, Cubango River launch site, -12.61700° 16.22133°, 1,727 m asl. Fig. 30. Adult male Ptychadena grandisonae from upper Cubango River. Photo by Werner Conradie. Amphib. Reptile Conserv. Elevation (m) =) So 3 Map 26. Distribution of Ptychadena bunoderma in Angola. Description: Medium sized Ptychadena; narrow body; ridges present on thigh; two and a half phalanges of the longest toe free of webbing. Dorsum light brown, with darker brown blotches; back of thighs with pale stripes; yellow infusion in groin. Only adult male measured 36.2 mm (PEM A12689). Habitat and natural history notes: This species was only encountered along the Cubango River in flooded grassland. Comments: The identification was based on the key provided by Poynton and Broadley (1985b) and needs genetic verification. Ptychadena upembae (Schmidt and Inger, 1959) Upemba Ridged Frog (Map 28) Material (1 specimen): PEM A14829, camp at side tributary (Luandai River) of the Luanguinga River, -13.70885° 21.26234°, 1,116 m asl. Description: Medium sized Ptychadena; elongated body; longitudinal dorsal ridges present; long toes; reduced webbing, with up to three phalanges of longest toe free of webbing. Dorsum brown to beige with black and brown spots; black facial mask, from snout through eye to forearm; broad light vertebral band present; thin light line on dorsal tibia; back of thighs with light and dark bands; ventrum light yellow. Only adult male measured 37.7 Elevation (m) Map 27. Distribution of Ptychadena grandisonae in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola Elevation (m) ra So oO Map 28. Distribution of Ptychadena upembae in Angola. mm (PEM A14829). Habitat and natural history notes: Found in grassland alongside a large river during the day. Comments: Identification was based on the key provided by Poynton and Broadley (1985b). Only a few records of this species exist for central and eastern Angola (Marques et al. 2018). Re-examination of material recorded by Conradie et al. (2016) of Pitychadena guibei also conform to this species. Ptychadena keilingi (Monard, 1937) Keiling’s Ridged Frog (Fig. 31; Map 29) Material (38 specimens): PEM A12428, Cutva River bridge on EN250, -11.98345° 17.72367°, 1,267m asl; PEM A12477-8, INBAC (no number x2), Cuanavale River source lake, -13.08537° 18.89098°, 1,360 m asl; PEM A12491, Dala River, near Samanunga village, -12.93170° 18.81458°, 1,363 m asl; PEM A12559-61, Cuito River source lake, -12.68935° 18.36012°, 1,431 m asl; PEM A12646, outlet of Cuito River source lake, -12.70453° 18.35445°, 1,429 m asl; PEM A12752-9, INBAC: WC- 4797, (no number x2), Cuando River source, -13.00346° 19.12751°, 1,353 m asl; PEM A12782, Quembo River, trap 1, -13.13592° 19.04417°, 1,369 m asl; PEM A12822— 4, Cutva River source, -12.66856° 18.35307°, 1,433 m Fig. 31. Adult male Ptychadena keilingi from Cuito River source. Photo by Werner Conradie. Amphib. Reptile Conserv. asl; PEM A12894—-6, INBAC: WC-4691, INBAC: (no number x1), Quembo River source lake, -13.13624° 19.04591°, 1,366 m asl; PEM A13818-19, en route to Cuemba Village, -11.96587° 17.76176°, 1,302 m asl; PEM A14694, Lungwebungu River camp, right bank past first oxbow, -12.57956° 18.67761°, 1,295 m asl; PEM A14736, Quembo River, walk back from small waterfall, -13.52988° 19.28340°, 1,242 m asl; PEM A14780, Quembo River bridge camp, trap 1, -13.52801° 19.28147°, 1,236 m asl; PEM A14781, INBAC: WC-6962, Quembo River right side tributary (Micongo River) past village, -13.51877° 19.28487°, 1,248 m asl; PEM A14894, wetland near old quarry east of Quemba, -12.16960° 18.22965°, 1,353 m asl. Additional specimens (2 specimens): SAIAB 204561 (2 specimens), Quembo River source lake, -13.13611° 19.04500°, 1,367 m asl. Description: Medium sized Ptychadena, elongated body; longitudinal dorsal ridges present; rostrum protruding well beyond lower jaw and forming a very pointed and elongated keratinized protrusion; long toes; reduced webbing, with up to four phalanges of longest toe free of webbing. Dorsum colorful: red infusions on the flanks; black facial mask, from snout through eye to forearm; broad light vertebral band present; thin light line on dorsal tibia; top of thighs with light and dark bands; ventrum immaculate, with black spots on chest in some specimens; black spot in front of arm that connects with lower jaw; lower jaw with spots; back of thighs with irregular longitudinal black and yellow stripes. Adult females (n = 19) varied from 24.6—39.9 (34.8) mm (largest female: PEM A12755); adult males (n= 14) varied from 25.3-31.5 (27.9) mm (largest male: PEM A14894). Habitat and natural history notes: Found in the flooded margins of rivers and lakes. Comments: Originally described from Dala, in north-eastern Angola (Laurent 1964). It is only known from northeastern Angola (Laurent 1964; Marques et al. 2018), western Zambia (Channing 2001), and southwestern DRC (Channing and Rodel 2019). This is the southernmost record for this species, and a new species record for southeastern Angola and the Okavango River basin. Elevation (m) 3 So oO I | | lee | 12 14 16 18 20 22 24 Map 29. Distribution of Ptychadena keilingi in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. Fig. 32. Adult male Ptychadena oxyrhynchus from Cuito town. Photo by Werner Conradie. Ptychadena oxyrhynchus (Smith, 1849) Sharp-nosed Ridged Frog (Fig. 32; Map 30) Material (6 specimens): PEM A12628, roadside ditch 10 km SW of Cuito town, -12.44815° 16.88118°, 1,742 m asl; PEM A12667, Campsite 1 below Cubango River rapids, west of Fundo village, -13.04483° 16.37520°, 1,585 m asl; PEM A12835-6, 31 km W of Menongue, Cueli River, -14.70511° 17.38014°, 1,392 m asl; PEM A14830, camp at side tributary (Luandai River) of the Luanguinga River, -13.70885° 21.26234°, 1,116 m asl. Additional material (1 specimen): SAIAB 204559 (1 specimen), half-way house road just outside Munhango village, pools in road. Description: Large and robust Ptychadena, sharp pointed snout; clear longitudinal dorsal ridges; extensive webbing, with one phalange on outer toe (5") free of webbing; large pointed subarticular tubercles on toes; small inner metatarsal tubercle; outer metatarsal tubercle absent; enlarged palmar tubercles; ventrum smooth. Dorsum light brown to gray, with scatted dark spots (PEM A12628 with unusual orange dorsum and yellow infusion in the outer thigh region); pale triangle on snout; lower jaw barred; back of thigh with irregular white blotches or spots that are sometimes fused; ventrum immaculate. Adult males (n = 5) varied from 44.1—51.9 (47.7) mm (largest male: PEM A12835). Fig. 33. Adult male Ptychadena porosissima from Lungwebungu River crossing. Photo by Werner Conradie. Amphib. Reptile Conserv. Elevation (m) ra) S S Map 30. Distribution of Ptychadena oxyrhynchus in Angola. Habitat and natural history notes: Only found in open dry savanna. Comments: This species has a wide distribution, from Senegal to South Africa (Channing and Roédel 2019). Found across most of Angola (Marques et al. 2018), associated with a wide variety of habitats. Smith (1849) gave the type locality as ‘Kaffirland and the region of Port Natal,’ which is situated along the east coast of South Africa. Material from Angola differs molecularly from the topotypical material and warrants further phylogenetic investigation (Hubler 2015). Ptychadena porosissima (Steindachner, 1867) Striped Ridged Frog (Fig. 33; Map 31) Material (17 = specimens): PEM A12712-6, Lungwebungu River camp bridge crossing, -12.58347° 18.66600°, 1,304 m asl; PEM A12901—2, PEM A12906, Quembo River source lake, -13.13624° 19.04591°, 1,366 m asl; PEM A13749-51, Lungwebungu River Campsite, -12.58319° 18.66573°, 1,284 m asl; PEM A13782-3, Lungwebungu River Trap 1, -12.58013° 18.66740°, 1,298 m asl; PEM A14708-9, INBAC: WC- 6750, Lungwebungu River camp, -12.58439° 18.66748°, 1,297 m asl; PEM A14782, Quembo River bridge camp, -13.52746° 19.28060°, 1,241 m asl. Description: Medium sized Ptychadena; sharp snout; large tympanum, Elevation (m) =} o Oo Map 31. Distribution of Ptychadena porosissima in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola almost equal to eye diameter; three phalanges of longest toe free of webbing; webbing reaches first phalange of the outer toe; large inner metatarsal tubercle; outer metatarsal tubercle inconspicuous or absent; distinct inner tarsal ridge; subarticular tubercles single and prominent. Dorsum brown, with scattered black spots along ridges; snout paler than body; vertebral stripe broad; dorsolateral ridge pale, elevated, and prominent; white anterior bars on lower jaw; dark facial mask from snout to front of arms; dorsal pale stripe on tibia; back of thighs with light spots forming longitudinal lines; ventrum immaculate, except for scattered faint black blotches between arms. In males, ventrum covered in small brown asperites: thumbs and nuptial pads swollen. Adult females (n = 7) varied from 34.8-46.2 (41.1) mm (largest female: PEM A12716); adult males (n = 7) varied from 34.9-40.0 (37.4) mm (largest male: PEM A14782). Habitat and natural history notes: Found in flooded grasslands or river margins. Comments: Found in sympatry with the closely related P. uzungwensis, but at lower abundance. The two species can be distinguished based on ventral asperites (present in P. porosissima versus absent in P. uzungwensis), dorsal coloration (striped in P. porosissima versus mostly spotted in P uzungwensis), and dorsal tibia line (always present in P. porosissima versus mostly absent in P. uzungwensis). Ptychadena subpunctata (Bocage, 1866) Speckled-bellied Ridged Frog (Map 32) Material (2 specimens): PEM A14874, Ninda River, -14.84018° 21.66556°, 1,081 m asl; PEM A14812, Luio River camp floodplains, -13.19711° 20.22194°, 1,181 m asl. Description: Large robust Ptychadena;, snout pointed; two phalanges of longest toe free of webbing: large tympanum; small elevated inner metatarsal tubercle; outer metatarsal tubercle absent. Dorsum brown, with large rounded dark brown spots; dorsal ridges pale; narrow vertebral stripe present; lower jaw barred; white spot below eye; back of thighs with two continuous black stripes below vent from knee to knee; white stripe on dorsal tibia; ventrum finely spotted. In males, the forearms and thumbs are swollen and nuptial pads are black. Adult Elevation (m) Map 32. Distribution of Ptychadena subpunctata in Angola. Amphib. Reptile Conserv. 46 males (n = 2) varied from 44.6-61.0 (53.1) mm (largest male: PEM A14874). Habitat and natural history notes: Only found at Zambezi River tributaries to the east and clearly absent from the source lake area. Comments: Although originally described from Angola, only a few records are known from the country (Marques et al. 2018). Ptychadena taenioscelis Laurent, 1954 Small Ridged Frog (Fig. 34; Map 33) Material (39 specimens, 2 tadpole lots): PEM A12479— 81, Cuanavale River source lake, -13.08537° 18.89098°, 1,360 m asl; PEM A12492, Dala River, near Samanunga village, -12.93169° 18.81458°, 1,371 m asl; PEM A12523, Cuiva River source, -12.66825° 18.35282°, 1,407 m asl; PEM A11609, PEM A12562-69, INBAC (no number x 4), Cuito River source lake, -12.68935° 18.36012°, 1,431 m asl; PEM A12663, Campsite 2 near old Cuvango Mission, -13.33451° 16.41280°, 1,534 m asl; PEM A12760, INBAC: WC-4750, INBAC (no number), Cuando River source, -13.00346° 19.12751°, 1,353 m asl; PEM A12767—-8, Cuando River source trap 1, -13.00393° 19.12808°, 1,351 m asl; PEM A12783-4, INBAC: WC-4600, Quembo River trap 1, -13.13592° 19.04417°, 1,369 m asl; PEM A12837, 31 km W of Menongue, Cueli River, -14.70511° 17.38014°, 1,392 m asl; PEM A12897-—8, INBAC (no number), Quembo Fig. 34. Adult female Ptychadena taenioscelis from Cuito River source. Photo by Werner Conradie. Elevation (m) Map 33. Distribution of Ptychadena taenioscelis in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. River source lake, -13.13624° 19.04591°, 1,367 m asl; PEM A13813, Lake Tchanssengwe, -12.41403° 18.64418°, 1,393 m asl; PEM A13820, en route to Cuemba Village, -11.96587° 17.76176°, 1,302 m asl; PEM A14092 (tadpoles), series of pans south of Tempué, -13.55719° 18.85519°, 1,315 m asl; PEM A14122 (tadpoles), Calua River source, 6 km SE of Curto River source, -12.73675° 18.39310°, 1,446 m asl; PEM A14691, Lungwebungu River camp, first oxbow on right side, -12.58117° 18.67106°, 1,294 m asl; PEM A14813- 6, INBAC: WC-7009, Luio River camp floodplains, -13.19711° 20.22194°, 1,181 masl. Additional material (2 specimens, 1 tadpole lot): SAIAB 204572 (2 specimens), Samaununga village, -12.93228° 18.81672°, 1,364 m asl; SAIAB 209102 (12 tadpoles), Calua lagoon, -12.73599° 18.39394°, 1,448 m asl. Description: Small Ptychadena; subarticular tubercles weakly developed; thin inner metatarsal tubercle; outer metatarsal tubercle absent or much reduced; three phalanges of longest toe free of webbing. Dorsum dark, with scattered black spots; broad vertebral band often present; white line above lip, from tip of snout to insertion of arm; bright green patch between eye and lip; dorsolateral ridge white; back of thighs with one continuous black stripe below vent from knee to knee; soles of feet, toes, forearms, and legs black: light line present on dorsal tibia; throat with scattered black spots; ventrum immaculate. Adult females (n = 21) varied from 21.5—33.2 (27.4) mm (largest female: PEM A12523); adult males (n = 9) varied from 23.1— 28.2 (26.5) mm (largest male: PEM A13813). Habitat and natural history notes: Found along river margins and floodplains. Comments: The close morphological relationship between P. pumilio and P. taenioscelis has been discussed in the past, and no clear concensus has yet been reached (Perret 1979; Poynton and Broadley 1987), although the former is mostly restricted to West Africa and the latter to Southern Africa (Channing and Rodel 2019). Bittencourt-Silva (2019) alludes to the close genetic relationship between P. pumilio and this species. Additional investigation is needed to validate its taxonomic status. Fig. 35. Adult male Ptychadena uzungwensis from Culua River source. Photo by Werner Conradie. Amphib. Reptile Conserv. Ptychadena uzungwensis (Loveridge, 1932) Udzungwa Ridged Frog (Fig. 35; Map 34) Material (56 specimens, 7 tadpole lots): PEM A12482-3, Cuanavale River source lake, -13.08537° 18.89098°, 1,360 m asl; PEM A12493, river crossing before Samanunga village, -12.93169° 18.81458°, 1,363 m asl; PEM A12570—2, Cuito River source lake, -12.68935° 18.36012°, 1,431 m asl; PEM A12609-11, PEM A14117 (tadpoles), Calua River source, 6 km SE of Cuito River source, -12.73675° 18.39310°, 1,445 m asl; PEM A12641, Cuanavale River, -13.29236° 18.96283°, 1,314 m asl; PEM A12717—8, Lungwebungu River camp bridge crossing, -12.58347° 18.66598°, 1,304 m asl; PEM A12761-—3, Cuando River Source, -13.00346° 19.12751°, 1,353 m asl; PEM A12769, Cuando River source, trap 1, -13.00393° 19.12808°, 1,351 m_ asl; PEM A12813-4, Cuito River source lake, -12.68727° 18.36067°, 1,424 m asl; PEM A12826, Quembo River, trap 3, -13.13073° 19.03725°, 1,453 m asl; PEM A12831-3, Culua River source, -12.73723° 18.39340°, 1,450 m asl; PEM A12838, 31 km W of Menongue, Cueli River, -14.70511° 17.38014°, 1,392 m asl; PEM A12899-900, PEM A12903-—5, Quembo River source lake, -13.13624° 19.04591°, 1,366 m asl; PEM A13752- 5, Lungwebungu River campsite, -12.58319° 18.66573°, 1,284 m asl; PEM A13814, Lake Tchanssengwe, -12.41403° 18.64418°, 1,393 masl; PEM A13816, John’s crossing before village, -12.20672° 18.06236°, 1,387 m asl; PEM A14083 (tadpoles), Longa rice paddies, Longa River, -14.56356° 18.44367°, 1,277 m asl; PEM A14084 (tadpoles), Cuanavale River source, -13.09033° 18.89396°, 1,359 m asl; PEM A14087 (tadpoles), 4 km upstream from Cuanavale River source, -13.05084° 18.89726°, 1,380 m asl; PEM A14093, (tadpoles) series of pans south of Tempué village, -13.55719° 18.85519°, 1,315 m asl; PEM A14095 (tadpoles), Dala River, near Samanunga village, -12.93169° 18.81458°, 1,363 m asl; PEM A14100 (tadpoles), 4 km downstream from Cuanavale River source camp, -13.11585° 18.90246°, 1,354 m asl; PEM A14692, Lungwebungu River camp, machamba (cultivated areas) on left side of river, f Elevation (m) Map 34. Distribution of Ptychadena uzungwensis in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola -12.58276° 18.66556°, 1,295 m asl; PEM A14695, Lungwebungu River camp, right bank past first oxbow, -12.58077° 18.67585°, 1,300 m asl; PEM A14710-11, Lungwebungu River camp, -12.58439° 18.66748°, 1,297 m asl; PEM A14719, PEM A14738, Quembo River bridge camp, -13.52746° 19.28060°, 1,241 m asl; PEM A14737, Quembo River bridge camp, trap 1, -13.52801° 19.28147°, 1,236 m asl; PEM A14760, Quembo River bridge camp, trap 2, -13.52816° 19.28067°, 1,240 m asl; PEM A14817, Luio River camp floodplains, -13.19711° 20.22194°, 1,181 m asl; PEM A14875, wetland south of Lake Hundo, -15.01099° 21.63608°, 1,100 m asl. Additional specimens (2 specimens): SAIAB 204568 (2 specimens), Quembo River source lake, -13.13611° 19.04500°, 1,367 m asl. Description: Medium sized Ptychadena, tympanum large, three-quarters the size of the eye; small inconspicuous metatarsal tubercle; large outer metatarsal tubercle; three phalanges of longest toe free of webbing; numerous irregular-sized palmar tubercles present; prominent longitudinal ridges; outer dorsolateral ridge broken anteriorly; no ventral asperites in males. Dorsum with scattered blotches, almost forming transverse bands; lower lip barred; vertebral stripe present or absent; snout distinctly marked, without pale triangle; back of thighs with large pale blotches, almost forming longitudinal stripes; light line on dorsal tibia often absent (present in PEM A12769, 12762, and 12900). Males with swollen and dark thumbs. Adult females (n = 25) varied from 27.5—43.7 (39.0) mm (largest females: PEM A12905, PEM A12831); adult males (n = 22) varied from 30.1—39.5 (35.0) mm (largest male: PEM A12611). Habitat and natural history notes: Commonly found in flooded grasslands adjacent to main rivers and source lakes. Comments: See P. porosissima account for information on sympatry between these two species. Pyxicephalidae Amietia angolensis (Bocage, 1866) Angola River Frog (Fig. 36; Map 35) Material (16 specimens): PEM A12627, roadside ditch 10 km SW of Cuito town, -12.44815° 16.88118°, wae Fig. 36. Adult male Amietia angolensis from west of Menongue. Photo by Werner Conradie. Amphib. Reptile Conserv. 1,742 m asl; PEM A12645, confluence of Cuito and Calua rivers, -12.74878° 18.35433°, 1,413 m asl; PEM A12653-7, INBAC: WC-5245, Kuvango Hydro Plant Site, -14.38775° 16.29365°, 1,441 m asl; PEM A12669, Campsite 1 below rapids, west of Fundo village, -13.04359° 16.37439°, 1,571 m asl; PEM A12671-3, INBAC: WC-5228, Camp 3, Malova Village, Mipanha River, -14.09140° 16.41476°, 1,553 masl; PEM A12834, 31 km W of Menongue, Cueli River, -14.70511° 17.38014°, 1,392 m asl; PEM A13785, Aquaculture farm outside Cuito, -12.43972° 16.89833°, 1,689 m asl; PEM A13786, Dam/Hydroplant on Rio Cuquema, -12.42556° 16.81856°, 1,640 m asl; PEM A14678, Villa Menongue, -14.63015° 17.63465°, 1373 m asl; bridge between Huambo and Cuito, -12.47056° 16.82389°, 1,643 m asl. Additional material (2 specimens): SAIAB 204011 (2 specimens), Cuchi River below bridge, -14.70272° 17.37864°, 1,393 m asl. Description: Large species; large tympanum, with an elevated ridge running from just behind the eye to arm; elevated paravertebral ridges continuous from behind eyes to groin, while the other ridges are broken (except in PEM A12834, where the dorsum is smooth); two and a half phalanges of longest toe free of webbing; small but conspicuous inner metatarsal tubercle; no outer metatarsal tubercle; well-developed subarticular tubercles; protruding eyes that are contained in outline of jaw viewed from below. Dorsum brown to dark green with scattered darker blotches; gular region dark with irregular white blotches that sometimes fuse to form lines; light vertebral stripe often present; dorsal tibia with dark crossbands; ventrum immaculate, except in two specimens (PEM A12654 and A12657) where the throat mottling extends onto the groin. In breeding males, the dorsum is yellowish, spiny with black-tipped asperites, and the nuptial pads on the thumbs are swollen. Adult males (n = 13) varied from 51.1—71.9 (63.9) mm (largest male: PEM A12671). Habitat and natural history notes: All material was found associated with the western tributaries of the Cubango River, where the substrate is rocky and the water is fast flowing. A single subadult specimen was found at the confluence of Elevation (m) 3 Oo So Map 35. Distribution of Amietia angolensis in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. Fig. 37. Adult male Zomopterna tuberculosa from west of Cuito town. Photo by Werner Conradie. the Calua and Cuito rivers. This species is absent from the rivers and floodplains of the main Okavango River basin, possibly due to the sandier substrate. Comments: Recent phylogenetic revisions of the genus led to the identification and description of numerous cryptic species (Channing and Baptista 2013; Larson et al. 2016; Channing et al. 2016). In the process, A. angolensis, which was once considered to have a wide distribution across most of southern African, was restricted to Angola and is now regarded as a country endemic (Channing and Baptista 2013; Channing et al. 2016). Tomopterna tuberculosa (Boulenger, 1882) Rough Sand Frog (Fig. 37; Map 36) Material (5 specimens): PEM A12632-3, roadside ditch 10 km SW of Cuito, -12.44815° 16.88118°, 1,743 m asl; PEM A12681, INBAC: WC-5235, Camp 3, Malova Village, Mipanha River, -14.09140° 16.41476°, 1,553 m asl; PEM A13792-—3, Dam/Hydroplant on Rio Cuquema, -12.42556° 16.81856°, 1,640 m asl. Description: Medium sized frog; short snout; tympanum clearly visible; elevated ridge from behind eye to arm; reduced webbing; inner metatarsal tubercle large; subarticular Fig. 38. Adult male Amnirana adiscifera stat. nov. from Cuando River source. Photo by James Harvey. Amphib. Reptile Conserv. Elevation (m) S oO So Map 36. Distribution of Zomopterna tuberculosa in Angola. tubercles single. Dorsum brown with near-symmetrical dark brown markings with thin white border; dark brown interorbital bar; no light vertebral line. Adult females (n = 2) measured 45.4-46.3 (45.8) mm (largest female: PEM A12681); adult males (n = 4) varied from 31.9- 36.5 (32.3) mm (largest male: PEM A12633). Habitat and natural history notes: This species was only found within the western side of the study area, associated with open and rocky habitats. Comments: This species occurs mostly in central and western Angola, with isolated records in the east (Marques et al. 2018). Elsewhere, it is known from northern Namibia and western Zambia to northern Tanzania and south to Zimbabwe (Channing and Rodel 2019). Ranidae Amnirana adiscifera (Schmidt and Inger, 1959) stat. nov. Green White-lipped Frog (Fig. 38; Map 37) Material (18 specimens, 5 tadpole lots): PEM A11599, Cuito River source lake, -12.68935° 18.36012°, 1,431 m asl; PEM A12447, Cuanavale River source lake, -13.08537° 18.89098°, 1,360 m asl; PEM A12726, Cuando River source, -13.00346° 19.12751°, 1,353 m Elevation (m) 5 o Oo Map 37. Distribution of Amnirana adiscifera stat. nov. in Angola. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola asl; PEM A12727-—8, Cuando River source, -13.00346° 19.12751°, 1,353 m asl; PEM A12764—6, Cuando River source, trap 1, -13.00393° 19.12808°, 1,351 m asl; PEM A12791, Cuanavale River source lake camp side, -13.09442° 18.89370°, 1,368 m asl; PEM A12798-9, Cuanavale River source lake opposite side, -13.08934° 18.89485°, 1,359 m asl; PEM A12842, west of Cuito town on Aludungo road, -14.64833° 16.97444°, 1,380 m asl; PEM A13736 (tadpoles), Comba River, -12.62442° 18.65159°, 1,299 m asl; PEM A13764, Lungwebungu River old oxbows, -12.58129° 18.67162°, 1,304 m asl: PEM A14082 (tadpoles), Luissinga River, -14.58899° 18.44367°, 1,311 m asl; PEM A14091 (tadpoles), Cuanavale River source, -13.09033° 18.89396°, 1,359 m asl; PEMA14099 (tadpoles), PEM A12485-—7, Dala River, near Samanga village, -12.93169° 18.81458°, 1,363 m asl; PEM A14121 (tadpoles), Culua River source, 6 km SE of Cuito River source, -12.73675° 18.39310°, 1,446 m asl; PEM A14679, Menongue, -14.63015° 17.63465°, 1,373 m asl; PEM A14716, Cuanavale River source lake, -13.09052° 18.89394°, 1,357 masl. Additional material (14 specimens, 4 tadpole lots): SAIAB 209135 (5 specimens), Kalilongue Dam _ inflow, -12.44722° 16.82428°, 1,429 m asl; SAIAB 209081 (1 specimen), swamp near Cuanavale River source, -13.10750° 18.86089°, 1,386 m asl; SAIAB 209080 (tadpoles), Cuito River source outlet, -12.70455° 18.35203°, 1,430 m asl; SAIAB 209088 (tadpoles), Calua River lagoon, -12.73599° 18.39394°, 1,448 m asl; SAIAB 209140 (tadpoles), Cuanavale River lake outlet, -13.09414° 18.89612°, 1,357 m asl; SAIAB 204541 (tadpoles), Cuanavale River source lake below fish fence, -13.09364° 18.89597°, 1,357 m asl; SAIAB 204567 (1 specimen), Quembo River source lake, -13.13611° 19.04500°, 1,363 m asl; SAIAB 204520 (2 specimens), SAIAB 204531 (1 specimen), SAIAB 204534 (1 specimen), Cuanavale River source lake, -13.08997° 18.89389°, 1,358 m asl: SAIAB 204484 (1 specimen), stream outflow about 2.6 km downstream of Cuando River source lake, -13.00317° 19.15153°, 1,333 m asl; SAIAB 204496 (2 specimens), river at Munhango lagoon, -12.17281° 18.54897°, 1,376 m asl. Description: Large ranid; large tympanum, nearly equal in size to eye; elevated upper lip ridge from snout tip to above arm: pair of dorsolateral ridges from eye to urostyle; some specimens have a small flap or ridge above the vent; smooth elevated subarticular tubercles on feet; small indistinct outer metatarsal tubercle, inner metatarsal tubercle present; pedal webbing formula: I (1), I w/e (1-2), TT i/e (1-3), TV (2-3), V (1); no dilated toe tips or terminal discs, no horizontal grooves present on toe tips; no external vocal sacs. All the adults collected were green and not the typical golden-brown coloration as illustrated in Du Preez and Carruthers (2009). Metamorphs retain dorsal and ventral spots, which fade during growth. Tadpoles are orange with black scattered dots. Males with a small nuptial pad on the thumb and enlarged glands on the upper arm. Adult Amphib. Reptile Conserv. females (n = 7) varied from 57.0—66.5 (61.6) mm (largest female: PEM A14716); adult males (n = 6) varied from 46.9-62.4 (55.0) mm (largest male: PEM A12799). Habitat and natural history notes: Males called in late afternoons and early evenings, from vegetation on the edge of open deep waters. Comments: Currently, five species of Amnirana have been recorded from Angola (Marques et al. 2018; Baptista et al. 2019): A. albolabris, A. darlingi, A. lemairei, A. lepus, and A. parkeriana. All except A. darlingi are forest-associated species, with toes that are dilated or have discs with horizontal grooves. The specimens found in our study do not exhibit dilated toes or terminal discs. Five specimens from Chitau that were initially referred to as Rana albolabris (see Schmidt 1936), were later described as Rana albolabris adiscifera by Schmidt and Inger (1959). Those authors only compared it to R. albolabris and R. a. lemairei, and differentiated it based on the condition of the toes (no dilated tips or discs), webbing (reduced versus extensive) and foot length (long versus short), respectively. Later, without much explanation, Laurent (1964) synonymised it with Rana darlingi |=Amnirana darlingi|, and this was followed by Poynton (1964) and Perret (1977). Jongsma et al. (2018) showed that eastern (Malawi) and western (Angolan) A. darlingi material differ significantly enough on a molecular level to warrant taxonomic re-evaluation. However, no taxonomic action has been undertaken pending more evidence. The type locality of A. darlingi is ‘Mazo6e and between Umtali [Mutare] and Marandellas [= Marondera], Mashonaland, Zimbabwe’ and represents the eastern material of Jongsma et al. (2018). The new series of specimens is in full morphological agreement with the description of R. a. adiscifera. However, no coloration details were provided in the original description or in Schmidt (1933). The newly collected material agrees in preserved coloration and morphology to photos of the holotype (https://collections-zoology. fieldmuseum.org/catalogue/1848393). The green Angolan form of A. darlingi is also in agreement with the green ‘undescribed Hillwood frog’ referred to by Channing (2001) and Channing and Rodel (2019). Recently, Ceriaco et al. (2016, 2018) also documented this green form of A. darlingi from central Angola. Based on the color differences, together with the typical form (green versus brown) and the genetic results by Jongsma et al. (2018), we here formally assign the green western material to Schmidt and Inger’s (1959) adiscifera and elevate it to full species, Amnirana adiscifera stat. nov. This species is widely distributed in central Angola and northwestern Zambia (see Channing 2001; Channing et al. 2013; Marques et al. 2018; Channing and Rodel 2019). Similar distribution patterns have been observed in Kassinula wittei, Kassina kuvangensis, as well as the snake Limnophis bicolor (Conradie et al. 2020a,b; this study). Amnirana can be divided into two morphotypes: those with short feet, dilated toes, or terminal discs and extensive webbing August 2023 | Volume 17 | Number 1 & 2 | e325 Conradie et al. (albolabris, amnicola, asperrima, fonensis, lemairei, lepus, occidentalis, and parkeriana), and those with long feet, reduced webbing, and no dilated toes or discs (adiscifera, darlingi, and galamensis). The former group is restricted to forests, while the latter prefers flooded grasslands in savanna (Poynton 1964). However, this pattern is not mirrored in the phylogenetic analysis, and indicates that adaptation from savanna to forest has occurred more than once (Jongsma et al. 2018). Discussion The material collected during this study contributes substantially to the knowledge of Angolan amphibians. Our findings have increased the number of documented amphibians for Angola by at least five species, three of which are potentially new (1.e., Hyperolius aff. bocagei, Kassinula cf. wittei, and Leptopelis sp.). A large collection of the green Amnirana ‘darlingi’ allowed us to reassess the taxonomic status of this material and led to both the reinstatement of Schmidt and Inger’s (1959) adiscifera and its elevation to full species as Amnirana adiscifera stat. nov. The rain frogs (Breviceps sp.) collected represent the first modern (post civil unrest, ~2002) material for Angola, and allowed us to explore their taxonomic status, which led to the description of a new endemic species, Breviceps ombelanonga (see Nielsen et al. 2020). Furthermore, the Kassinula wittei specimens represent the first country records, a range extension of over 500 km, and further allowed us to revisit the taxonomic relationships of the group relative to other hyperoliids (Conradie et al. 2020b). Conradie et al. (2020b) and Nielsen et al. (2020) highlighted the relevance of these new collections on the national and continental levels, as well as the necessity for further studies to document the diversity and distribution of Angolan biodiversity. Some of the species recorded here at the headwaters of the Okavango (e.g., Amnirana adiscifera stat. nov., Kassinula wittei, and Kassina kuvangensis) also occur in northwestern Zambia and eastern DRC (Poynton and Broadley 1985a,b, 1987, 1988; Channing 2001). Cei (1977) suggested that the amphibian fauna located south of the main Congo rivers to the Cubango rivers should have close affinities, given the lack of natural barriers. The limited genetic differences between Kassinula wittei from Angola and DRC (Neéas et al. 2022) provided support for this hypothesis, and this may also be reflected in other taxa. On a more regional level, we increased the number of amphibian species recorded from the Cuito, Cuanavale, and Cuando rivers in Angola by Conradie et al. (2016) by nine species, and documented an additional 42 species associated with the Angolan Zambezi River basin. Ten of the species recorded here represent the southeasternmost records for Angola, some of which are also new records for the larger Okavango River basin (1.e., Arthroleptis Amphib. Reptile Conserv. 51 stenodactylus, A. xenochirus, Breviceps ombelanonga, Hyperolius raymondi, Kassinula wittei, Ptychadena bunoderma, P. keilingi, Leptopelis sp., and Hyperolius aff. bocagei). In total, 125 nights, representing 240 total trapping nights, were spent surveying the region during both wet and dry seasons, without adding any additional species to the checklist. Thus, we now consider the upper catchments of the Cuito, Cuanavale, Cuando, and Lungwebungu rivers to be well surveyed for amphibians. However, the following species are expected to occur eastward to the Zambian border (compare to Broadley 1971; Pietersen et al. 2017; Bittencourt-Silva 2019), based the availability of appropriate habitat: Breviceps adspersus, Breviceps poweri, Chiromantis xerampelina, Hildebrandtia ornata, Hemisus marmoratus, Phrynomantis bifasciatus, Phrynomantis affinis, Poyntonophrynus fenoulheti, Poyntonophrynus kavangensis, Ptychadena mapacha, Pyxicephalus adspersus, and Tomopterna cryptotis. The headwaters of the Okavango River basin, with peat lakes and extensive floodplains, harbor a high diversity of Hyperoliidae (three genera, comprising 12 species) and Ptychadenidae (nine species), compared to the more terrestrial families such as Bufonidae. Many of the species are regarded as local endemics to the headwaters (1.e., Hyperolius raymondi, Hyperolius aff. bocagei, Kassina kuvangensis, Kassinula_ wittei, and Ptychadena keilingi). The increased knowledge coupled with the presence of habitat specialist species will contribute to better conservation planning in these regions and the larger Okavango River basin. Acknowledgments.—We thank the Wild Bird Trust, which administers the National Geographic Okavango Wilderness Project (2016-2019 National Geographic Society grant). Material was collected and exported under the following export permits issued by the Angolan Ministry of Environment Institute of Biodiversity (MINAMB): 31/GGPCC/2016, 89/INBAC. MINAMB/2017, 002/GGPTBOK/18, and 151/INBAC/ MINAMB/2019. Ethical clearance for this study was obtained from the Port Elizabeth Museum (Bayworld) ethics committee (Ethical Clearance no. 2013 and 2017-2). We would also like to acknowledge the use of infrastructure and equipment provided by the NRF- SAIAB Aquatic Genomics Research Platform, and the funding channeled through the NRF-SAIAB Institutional Support System for generating barcodes. This project is endorsed and supported by the Governors of Cuando Cubango, Bié and Moxico provinces. Various colleagues are thanked for collecting herpetological material during their fieldwork, including Alexander Rebelo, Timdteo Julio, Gotz Neef, Roger Bills, Paul Skelton, Maans Booysens, and Kerllen Costa. WC thanks the Eastern Cape Province Department of Sport, Recreation, Arts, and Culture (DSRAC) and Port Elizabeth Museum (Bayworld) for granting special leave to take part in August 2023 | Volume 17 | Number 1 & 2 | e325 Amphibians of the Okavango Delta headwater area in Angola these surveys. We thank Enviro-Insight for constructing and donating the drift fences required for the trapping exercise. NLB is currently supported by BIOPOLIS 2022-19. Literature Cited Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ. 1990. Basic local alignment search tool. Journal of Molecular Biology 215: 403-410. Angel MF. 1924. 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Perret JL. 1977. Les Hylarana (Amphibiens, Ranidés) du Cameroun. Revue Suisse de Zoologie 84: 841-868. Perret JL. 1979. Remarques et mise au point sur quelques especes de Ptychadena (Amphibia, Ranidae). Bulletin de la Société des Sciences Naturelles de Neuchdtel 102: 5-21. Pickersgill M. 2007. Frog Search. Results of Expeditions to Southern and Eastern Africa from 1993-1999. Frankfurt Contributions to Natural History 28. Edition Chimaira, Frankfurt, Germany. 574 p. Pietersen DW, Pietersen EW, Conradie W. 2017. Preliminary herpetological survey of the Ngonye Falls and surrounding regions in south-western Zambia. Amphibian & Reptile Conservation 11(1) [Special Section]: 24—43 (e148). Poynton JC. 1964. The Amphibia of Southern Africa: a faunal study. Annals of the Natal Museum 17: 1-334. Poynton JC, Broadley DG. 1985a. Amphibia Zambesiaca 1. Scolecomorphidae, Pipidae, Microhylidae, Hemisidae, Arthroleptidae. Annals of the Natal Museum 26(2): 503-553. Poynton JC, Broadley DG. 1985b. Amphibia Zambesiaca 2. Ranidae. Annals of the Natal Museum 27(1): 115— 181. Poynton JC, Broadley DG. 1987. Amphibia Zambesiaca 3. Rhacophoridae and Hyperoliidae. Annals of the Natal Museum 28(1): 161-229. Poynton JC, Broadley DG. 1988. Amphibia Zambesiaca 4. Bufonidae. Annals of the Natal Museum 29(2): 447-490. Poynton JC, Broadley DG. 1988. Amphibia Zambesiaca 5. Zoogeography. Annals of the Natal Museum 32(1): 221-277. Poynton JC, Loader SP, Conradie W, Rodel M-O, Liedtke C. 2016. Designation and description of a neotype of Sclerophrys maculata (Hallowell, 1854), and reinstatement of S. pusilla (Mertens, 1937) (Amphibia: Anura: Bufonidae). Zootaxa 4089: 73-94. Roédel M-O. 2000. Herpetofauna of West Africa, Volume I; Amphibians of the West African Savanna. Edition Chimaira, Frankfurt, Germany. 322 p. Ruas C. 1996. Contribui¢ao para o conhecimento da fauna de batraquios de Angola Parte I: Familias Pipidae, Bufonidae, Microhylidae, Ranidae, Hemisidae e Amphib. Reptile Conserv. Arthleptidae. Garcia de Orta. Series Zoology (Lisboa) 21(1): 19-41. Ruas C. 2002. Batraquios de Angola em colec¢ao no Centro de Zoologia. Garcia de Orta. Series Zoology (Lisboa) 24(1—2): 139-146. Schiotz A. 1999. Treefrogs of Africa. Editions Chimaira, Frankfurt am Main, Germany. 350 p. Schmidt KP. 1936. The amphibians of the Pulitzer- Angola Expedition. Annals of Carengie Museum 25: 127-133. Schmidt KP, Inger RF. 1959. Amphibians exclusive of the genera Afrixalus and Hyperolius. Exploration du Parc National de I’ Upemba. Mission G.F- de Witte, en collaboration avec W. Adam, A. Janssens, L. van Meel et R. Verheyen (1946-1949) 56: 1-264. Smith A. 1849. ///ustrations of the Zoology of South Africa; Consisting Chiefly of Figures and Descriptions of the Objects of Natural History Collected during an Expedition into the Interior of South Africa, in the Years 1834, 1835, and 1836. Volume III. Reptilia, Part 28. Smith, Elder, and Company, London, United Kingdom. 94 p. Stanley EL, Ceriaco LMP, Bandeira S, Valerio H, Bates MF, Branch WR. 2016. A review of Cordylus machadoi (Squamata: Cordylidae) in southwestern Angola, with the description of a new species from the Pro-Namib desert. Zootaxa 4061(3): 201-226. Telford N, Alexander GJ, Becker FS, Conradie W, Jordaan A, Kemp L, Le Grange A, Rebelo AR, Strauss S, Taft JM, et al. 2022. Extensions to the known geographic distributions of reptiles in the Karoo, South Africa. Herpetological Conservation and Biology 17(1): 145-154. Wagner P, Butler BO, Ceriaco LM, Bauer AM. 2021. A new species of the Acanthocercus atricollis (Smith, 1849) complex (Squamata, Agamidae). Salamandra 57: 449-463. Zimkus BM, Schick S. 2010. Light at the end of the tunnel: insights into the molecular systematics of East African Puddle Frogs (Anura: Phrynobatrachidae). Systematics and Biodiversity 8: 39-47. Zimkus BM, Rodel MO, Hillers A. 2010. Complex patterns of continental speciation: molecular phylogenetics and biogeography of sub-Saharan Puddle Frogs (Phrynobatrachus). Molecular Phylogenetics and Evolution 55(3): 883—900. August 2023 | Volume 17 | Number 1 & 2 | e325 Amphib. Reptile Conserv. Amphibians of the Okavango Delta headwater area in Angola Werner Conradie has a Masters in Environmental Science (M.Env.Sc.) and 18 years of experience working with the southern African herpetofauna. His main research interests focus on the taxonomy, conservation, and ecology of amphibians and reptiles. Werner has published numerous principal and collaborative scientific papers, and has served on many conservation and scientific panels, including the Southern African Reptile and Amphibian Relisting Committees. He has undertaken research expeditions to many African countries, including Angola, Botswana, Lesotho, Malawi, Mozambique, Namibia, South Africa, Zambia, and Zimbabwe. Werner is currently the Curator of Herpetology at the Port Elizabeth Museum (Bayworld), South Africa. Chad Keates is currently a Post-doctoral Fellow at the Port Elizabeth Museum, funded by research funds through Nelson Mandela University (Port Elizabeth, South Africa). Having recently completed his Ph.D. in Zoology, Chad’s research focuses on the African herpetofauna and its evolutionary and ecological structuring. In Chad’s short professional career, he has published several principal and collaborative peer-reviewed scientific papers and book chapters. As a strong advocate for reptile and amphibian awareness, Chad regularly conducts walks, talks, and presentations; and he has produced numerous popular scientific outputs on the subject. He has undertaken numerous expeditions in various African countries, such as Angola, Zambia, and South Africa, with a variety of both professional and scientific organizations. Luke Verbursgt is a consulting herpetologist in South Africa with over 19 years of herpetofauna survey experience across 23 African countries (Angola, Botswana, Cameroon, Democratic Republic of Congo, Ghana, Ivory Coast, Kenya, Lesotho, Liberia, Namibia, Madagascar, Malawi, Mali, Morocco, Mozambique, Republic of Guinea, SAo Tomé and Principe, Sierra Leone, South Africa, Swaziland, Tanzania, Uganda, and Zimbabwe). He is a co-owner of Enviro-Insight (Pretoria, South Africa), holds an M.Sc. in Zoology from the University of Pretoria, and is a registered scientific professional with the South African Council for Natural Scientific Professions (SACNASP). Luke has published more than 30 scientific articles, which include the descriptions of several new African herpetofauna species, and he is a co-author of the book Snakes and other Reptiles of Zambia and Malawi (Struik Random House Publishers, Cape Town, South Africa). He is also an extraordinary lecturer in the Department of Zoology & Entomology at the University of Pretoria. Ninda Baptista is an Angolan biologist with an M.Sc. degree in Conservation Biology from the University of Lisbon (Portugal). She is currently pursuing a Ph.D. in Biodiversity, Genetics, and Evolution at the University of Porto (Portugal) that addresses the diversity of Angolan amphibians. Over the last 13 years, she has worked on research, in-situ conservation projects, and environmental consulting in Angola, including priority areas for conservation along the Angolan escarpment and highlands. She has conducted herpetological surveys throughout the country, and created a herpetological collection (Coleccaéo Herpetologica do Lubango) that is currently deposited in Instituto Superior de Ciéncias da Educacado da Huila (ISCED — Huila) in Angola. Ninda is an author of various scientific papers and book chapters on Angolan herpetology and ornithology. She also works on scientific outreach, producing magazine articles, books for children, and posters about the country’s biodiversity in collaboration with Fundagéo Kissama (Luanda, Angola). James Harvey lives in South Africa and works as an independent herpetologist, ecological researcher, and consultant. He holds degrees in Zoology, Hydrology, and Environmental Management, and has performed herpetological fieldwork widely, primarily within Africa, in such places as South Africa, Botswana, Zimbabwe, Angola, Malawi, Kenya, Mali, Democratic Republic of Congo, Madagascar, and Vietnam. His interests are diverse but center on the taxonomy, ecology, and conservation of herpetofauna and other biodiversity. James has contributed to conservation assessments, workshops, and Red Data publications for reptiles, amphibians, mammals, and plants for the southern and eastern African regions. James regularly attends herpetological conferences, has published several scientific papers, and has contributed to a number of herpetological publications as an author. 56 August 2023 | Volume 17 | Number 1 & 2 | e325 The herpetofauna of the Baja California Peninsula endemic to the southern island of Isla Todos Santos, BC [Baja California]” (Grismer, 2002: 281). This individual was found on Isla Todos Santos Sur, Baja California, in the municipality of Ensenada. Isla Todos Santos Sur is one of a pair of islands lying just over 19 km off the coast of Ensenada in northern Baja California. Grismer (2002: 281) indicated that this snake “appears to be restricted to the rocky interior area of Isla Sur” and that it feeds on the lizards Sceloporus occidentalis and Plestiodon skiltonianus. Some authors have considered this kingsnake as a subspecies of Lampropeltis zonata (e.g., Heimes 2016), but our position on the validity of subspecies (see Materials and Methods) supports our recognition of this snake as a distinct species, a position that is also recognized by Grismer (2002) and on the Mesoamerican Herpetology website (http://www.mesoamericanherpetology.com; accessed 11 February 2023). Wilson et al. (2013a) calculated the EVS of this snake as 20, placing it in the highest vulnerability category. The IUCN does not recognize the taxonomy of this snake, and SEMARNAT considers it as Threatened (A). Photo by Jorge H. Valdez-Villavicencio. Official journal website: amphibian-reptile-conservation.org Amphibian & Reptile Conservation 17(1 & 2) [General Section]: 57-142 (e326). The herpetofauna of the Baja California Peninsula and its adjacent islands, Mexico: composition, distribution, and conservation status ‘Anny Peralta-Garcia, ‘Jorge H. Valdez-Villavicencio, 7Lydia Allison Fucsko, 3Bradford D. Hollingsworth, *Jerry D. Johnson, ‘Vicente Mata-Silva, Arturo Rocha, 5Dominic L. DeSantis, ‘Louis W. Porras, and ’*Larry David Wilson 'Conservacion de Fauna del Noroeste, A.C., Ensenada, Baja California 22897, MEXICO *Department of Humanities and Social Sciences, Swinburne University of Technology, Melbourne, Victoria, AUSTRALIA *Department of Herpetology, San Diego Natural History Museum, San Diego, California, 92101, USA *Department of Biological Sciences, The University of Texas at El Paso, El Paso, Texas 79968-0500, USA *Department of Biological & Environmental Sciences, Georgia College & State University, Milledgeville, Georgia 31061, USA °7705 Wyatt Earp Avenue, Eagle Mountain, Utah 84005, USA ‘Centro Zamorano de Biodiversidad, Escuela Agricola Panamericana Zamorano, Departamento de Francisco Morazdn, HONDURAS °1350 Pelican Court, Homestead, Florida 33035-1031, USA Abstract.—The herpetofauna of the Baja California Peninsula, Mexico, consists of 172 species, including 18 anurans, three salamanders, 140 squamates, and 11 turtles. Among the 10 recognized geographic regions, the total number of herpetofaunal species ranges from a low of 27 in the Baja California Coniferous Forest Region to 84 in the Gulf Islands Region. The individual species occupy from one to 10 regions (xX = 3.3). The Gulf Islands Region is of the greatest significant conservation importance, inasmuch as it contains the largest overall number of species (84), the largest number of single-region species (39), and the greatest number of peninsular endemics (50). Asimilarity dendrogram based on the Unweighted Pair Group Method with Arithmetic Averages (UPGMA) indicates that the two most closely related regions are the Central Gulf Coast Region and the Arid Tropical Region, while the most distantly related region is the Baja California Coniferous Forest Region. Among the distributional categories, the greatest number of species are the non-endemics (81 of 172, or 47.1%), followed by the peninsular endemics (77, 44.8%), and finally, the non-natives (14, 8.1%). The principal environmental threats to the herpetofauna of the Baja California Peninsula are land conversion and habitat loss, water diversion and overuse, invasive species, livestock grazing, illegal trade, off-road activities, infectious diseases, and climate change. We assessed the conservation status of the native species by employing the SEMARNAT, IUCN, and EVS systems. Of the 158 native species, 85 are included in NOM-059-SEMARNAT, 15 are in the IUCN Threatened categories, and 76 have high EVS values. Two Relative Herpetofaunal Priority (RHP) methods were used to identify the rank order significance of the 10 geographic regions, and the highest ranks for both methods were obtained for the Gulf Islands Region. Thirty protected areas are located in the Baja California Peninsula, and they comprise almost one-half of the total area. All but eight of the 158 native herpetofaunal species are represented within the system of protected areas. Few herpetofaunal surveys of the protected areas have been completed thus far, so this is a major conservation goal for the future. Keywords: Anurans, caudates, conservation status, physiographic regions, protected areas, protection recom- mendations, squamates, turtles Resumen.—La herpetofauna de la Peninsula de Baja California, México, consta de 172 especies, incluyendo 18 anuros, tres salamandras, 140 escamosos y 11 tortugas. Entre las 10 regiones geograficas reconocidas, el numero total de especies de herpetofauna varia desde un minimo de 27 en la Region del Bosque de Coniferas de Baja California hasta 84 en la Region de las Islas del Golfo. Las especies individuales ocupan de una a 10 regiones (xX = 3.3). La Region de las Islas del Golfo es de gran importancia para la conservacion, ya que contiene el mayor numero total de especies (84), el mayor numero de especies de una sola region (39) y el mayor numero de endémicas peninsulares (50). Un dendrograma de similitud basado en el método de grupos de pares no ponderados con promedios aritmeticos (UPGMA) indica que las dos regiones mas estrechamente relacionadas son la Region de la Costa Central del Golfo y la Region Tropical Arida. La region mas lejanamente relacionada es la Region del Bosque de Coniferas de Baja California. Entre las categorias de distribucion el mayor numero de especies son las no endemicas (81 de 172 0 47.1%), seguidas de las endémicas peninsulares (77 0 44.8%) y, por ultimo, las no nativas (14 0 8.1%). Las principales amenazas ambientales para la herpetofauna de la peninsula de Baja California son la conversion de tierras y la perdida de habitat, el desvio y uso excesivo Correspondence. = annyperalta@faunadelnoroeste.org (APG), jh valdez@yahoo.com.mx (JHVV), lyvdiafucsko@gmail.com (LAF), bhollingsworth@sdnhm.org (BDH), jjohnson@utep.edu (JDJ), vmata@utep.edu (VMS), arocha3@miners.utep.edu (AR), dominic.desantis@gcsu.edu (DLD), empub@msn.com (LWP), bufodoc@aol.com (LDW) Amphib. Reptile Conserv. 57 November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula de agua, las especies invasoras, el pastoreo de ganado, el comercio ilegal, las actividades todoterreno, las enfermedades infecciosas y el cambio climatico. Evaluamos el estado de conservacion de las especies nativas empleando los sistemas de SEMARNAT, UICN y EVS. De las 158 especies nativas, 85 estan incluidas en la NOM-059-SEMARNAT, 15 en las categorias Amenazadas de la UICN y 76 presentaron valores elevados de EVS. Mediante el uso de los dos métodos de prioridad herpetofaunistica relativa (RHP) para identificar la importancia del orden de rango de las 10 regiones geograficas, se obtuvieron los dos rangos mas altos para la region de las Islas del Golfo. Treinta areas protegidas se encuentran en la Peninsula de Baja California y comprenden casi la mitad del area total. Todas menos ocho de las 158 especies nativas de herpetofauna estan representadas dentro del sistema de areas protegidas. Se han completado pocos estudios de herpetofauna para las areas protegidas, por lo que este es un objetivo de conservacion importante para el futuro. Palabras Claves: Anuros, caudados, escamosos, tortugas, regiones fisiograficas, areas protegidas, estatus de conservacion, recomendaciones de proteccion Citation: Peralta-Garcia A, Valdez-Villavicencio JH, Fucsko LA, Hollingsworth BD, Johnson JD, Mata-Silva V, Rocha A, DeSantis DL, Porras LW, and Wilson LD. 2023. The herpetofauna of the Baja California Peninsula and its adjacent islands, Mexico: composition, distribution, and conservation status. Amphibian & Reptile Conservation 17(1&2): 57-142 (e326). Copyright: © 2023 Peralta-Garcia et al. This is an open access article distributed under the terms of the Creative Commons Attribution License [Attribution 4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org. Accepted: 2 August 2023; Published: 19 November 2023 “ .. Baja California possesses a wild, intangible allure. An innate sense of adventure, heightened by a hint of danger and the unknown, beckons many explorers off the highway into Baja‘’s harsh, poorly understood interior. Baja California’s jagged, snow-covered peaks, volcanic badlands, parched deserts with relentless summer temperatures, and arid, uninhabited desert islands have been reluctant to give up their secrets. There is still much knowledge to be attained and many personal challenges to be met.” L. Lee Grismer (2002) Introduction One of the most distinctive features of the physiography of Mexico is the Baja California Peninsula, a finger- like extension of land flanked by small islands, which comprises two states of the 32 federal entities in the country, 1.e., Baja California to the north and Baja California Sur to the south. These two states are separated from one another at latitude 28° N, just slightly to the north of Punta Eugenia, the northwestern point of the Peninsula de Vizcaino (Grismer 2002). A succinct description of the physiography of the peninsula is provided in the now-classic volume on the herpetofauna authored by L. Lee Grismer (2002), in which the introduction (pg. 3) includes the following quote: “Today Baja California is a thin northwest to southeast-tending peninsula nearly 1,300 km long. It is situated between 32° 30’ N latitude and 117° W longitude at its northwestern corner and [between] 23° N and 110° W at its southern tip. Its width ranges from approximately 240 km along the U.S. —Mexico border to less than 30 km at the Isthmus of La Paz. It is separated from the state of Sonora by the Rio Colorado in the north and from the rest of Sonora and mainland Mexico by the Gulf of Amphib. Reptile Conserv. California, approximately 160 km wide. The area of Baja California is approximately 143,400 km/?, and its coastline is approximately 3,300 km long. Associated with the coastline are forty-five major islands, each at least 1.3 km? in area. Several smaller islands are also associated with Baja California, and an additional 10 or so major islands are principally associated with the Mexican states of Sonora and Sinaloa... [which are not considered in this paper].” More details on the physiography of this peninsula are provided in the section below entitled “Physiography and Climate.” With an area of 71,450 km”, the state of Baja California is the 12" largest in Mexico and the 19" most densely populated. The corresponding data for the state of Baja California Sur are 73,909 km7, the 9" largest, and the 32" most densely populated (http://inegi.org.mx; accessed 8 June 2023). Given the relative geographic isolation of the peninsula from the mainland of both the United States and Mexico, this area is expected to be characterized by a significant degree of endemism, especially since the mainland of the peninsula is flanked by a large number of variously sized islands. Also, given the limited range of these endemic species, they are expected to be subjected to the usual range of anthropogenic threats. Thus, the purpose of this paper is to examine these aspects as they relate to the interesting herpetofauna of this offset region of Mexico. Materials and Methods Our Taxonomic Position In this paper, we follow the same taxonomic position as explained in previous works on other portions of Mesoamerica (Johnson et al. 2015; Mata-Silva et al. 2015; Teran-Juarez et al. 2016; Woolrich-Pifia et al. November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. 2016; Nevarez-de los Reyes et al. 2016; Cruz-Saenz et al. 2017; Gonzalez-Sanchez et al. 2017; Woolrich-Pifia et al. 2017; Lazcano et al. 2019; Ramirez-Bautista et al. 2020; Torres-Hernandez et al. 2021; Cruz Elizalde et al. 2022). Johnson et al. (2015) can be consulted for a detailed statement of this position, with special reference to the subspecies concept. System for Determining Distributional Status The system developed by Alvarado-Diaz et al. (2013) for the herpetofauna of Michoacan was employed to ascertain the distributional status of members of the herpetofauna of the Baja California Peninsula, which consists of the following four categories: SE, endemic to the Baja California Peninsula; CE, endemic to Mexico; NE, not endemic to Mexico; and NN, non-native in Mexico. Systems for Determining Conservation Status The following three systems were used to determine the conservation status of the 158 native species of amphibians and reptiles in the Baja California Peninsula; SEMARNAT, IUCN, and EVS. The SEMARNAT system, established by the Secretaria de Medio Ambiente y Recursos Naturales, lists only the threatened species in the NOM 059-SEMARNAT-2010 (SEMARNAT 2010, 2019) under three categories: Endangered (P), Threatened (A), and Subject to Special Protection (Pr). For species not included on that list, we used the designation “No Status;” however, we acknowledge that the SEMARNAT list is not meant to include non-threatened species. For species included on that list for which taxonomy has changed, we maintain the conservation status of the previous taxonomic entity, following section six of NOM-059-SEMARNAT-2010 (SEMARANT 2010). The IUCN system (https://www.iucnredlist.org) is utilized widely for assessing the conservation status of species on a global scale. The categories include Extinct (EX), Extinct in the Wild (EW), Critically Endangered (CR), Endangered (EN), Vulnerable (VU), Near Threatened (NT), Least Concern (LC), Data Deficient (DD), and Not Evaluated (NE). Collectively, the three categories of Critically Endangered, Endangered, and Vulnerable are termed the “threat categories” to distinguish them from the other six. The EVS system was applied here for the 158 native species, following Wilson et al. (2013a,b). A re- evaluation was conducted because of taxonomic changes that have transpired since their original EVS values were reported, as well as regional differences in the degree of human persecution in mainland Mexico, as required for criterion C. The EVS measure was not designed for use with marine species (e.g., marine turtles and sea snakes), and 1s generally not applied to non-native species. Amphib. Reptile Conserv. The Mexican Conservation Series The Mexican Conservation Series (MCS) was initiated in 2013, with a study of the herpetofauna of Michoacan (Alvarado-Diaz et al. 2013) that was published as part of a set of five papers designated as the Special Mexico Issue in Amphibian & Reptile Conservation. The basic format of the entries in the MCS was established in that paper, i1.e., examining the composition, physiographic distribution, and conservation status of the herpetofauna of a given Mexican state or group of states. Two years later, the MCS resumed with a paper on the herpetofauna of Oaxaca (Mata-Silva et al. 2015). That year, Johnson et al. (2015) authored a paper on the herpetofauna of Chiapas, and three entries in the MCS appeared the next year, covering Tamaulipas (Teran-Juarez et al. 2016), Nayarit (Woolrich-Pifia et al. 2016), and Nuevo Leon (Nevarez-de los Reyes et al. 2016). The following year three additional entries appeared, covering Jalisco (Cruz-Saenz et al. 2017), the Mexican Yucatan Peninsula (Gonzalez-Sanchez et al. 2017), and Puebla (Woolrich- Pifa et al. 2017). Subsequently, similar articles on Coahuila (Lazcano et al. 2019), Hidalgo (Ramirez- Bautista et al. 2020), and Veracruz (Torres-Hernandez et al. 2021) were published. Last year, articles on Querétaro (Cruz Elizalde et al. 2022), Tabasco (Barragan- Vazquez et al. 2022), and Guanajuato (Leyte-Manrique et al. 2022) appeared. Thus, this article on the herpetofauna of the Baja California Peninsula is the 16" entry in this series. Geography and Climate Geographic Regions The formation of the Baja California Peninsula in northwestern Mexico originated from a complex interaction of plate tectonics, which resulted in the formation of the Gulf of California. There are a large number of climatic variables and a varied topography along the peninsula (Shreve and Wiggins 1964; Grismer 2002; Hollingsworth et al. 2015). The interactions among these climatic and topographic variables have given rise to several distinct physiographic regions. These characteristics also resulted in the formation of different phytogeographical regions, which can be distinguished by the different types of vegetation (Fig. 1). This situation is linked to the “double ecological polarity” that occurs on the peninsula (Gonzalez- Abraham et al. 2010). First, a north-to-south gradient with a temperate climate in the northwest and a tropical climate in the south contains an extensive arid transition region between the two. Then, an east-to-west gradient has resulted from the combination of a mountain range distributed intermittently along the peninsula and the influence of two different marine water masses which are cold in the Pacific and warm in the Gulf of California (Gonzalez-Abraham et al. 2010). November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula 118°0'0"W 116°0'O"W 32°0'0"N 30°0'0"N 28°0'0"N Physiographic regions "> California ~ Baja California Coniferous Forest ~~ Lower Colorado Valley \— Vizcaino Central Gulf Coast )) Magdalena Arid Tropical MY Sierra La Laguna MW Pacific Islands > Gulf Islands 26°0'0"N 24°0'0"N 118°0'0"W 116°0'0"W 114°0'O"W 114°0'0"W 112°0'0"W 110°0'0"W 32°0'0"N 30°0'0"N 28°0'0"N 26°0'0"N 24°0'0"N 112°0'0"W 110°0'O"W Fig. 1. Geographic regions of the Baja California Peninsula, Mexico. The distribution of the herpetofauna of the Baja California Peninsula coincides with the phytogeographic regions (Grismer 1994, 2002). As indicators of the natural biotic provinces, they likely are influenced by the same environmental characteristics that limit the distributions of amphibians and reptiles (Grismer 2002). The following 10 regions have been identified: (1) California; (2) Coniferous Forest; (3) Lower Colorado Desert; (4) Vizcaino Desert; (5) Central Gulf Coast Desert; (6) Magdalena; (7) Sierra La Laguna; (8) Arid Tropical; (9) Pacific Islands; and (10) Gulf Islands. Shreve and Wiggins (1964), Wiggins (1980), and Grismer (2002) defined these regions, and they are briefly described below. Amphib. Reptile Conserv. 60 California Region. Located in the northwestern quarter of the peninsula, this region (Fig. 2) extends 275 km from the border with the United States to the vicinity of El Rosario along the Pacific coast, where it slowly intergrades into the Vizcaino Desert Region to the south (Hollingsworth et al. 2015). During the Pleistocene, the California Region likely extended farther southward onto the peninsula. However, successive periods of aridification replaced these communities with deserts that include remnants of the California Region flora (Van Devender 1990). Today, the California Region is confined to the northern state of Baja California. To the east, the region extends up the Sierra Juarez and Sierra San Pedro Martir to an elevation of 2,000 m at the start November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. (3,100 m), is seen in the background. Photo by Felipe Leon. of the pine belt, with a mixed chaparral-coniferous forest zone found at elevations between 1,500 and 2,000 m (Delgadillo 2004). Farther to the east, the California Region comes into contact and intermixes with the Lower Colorado Desert in low-elevation passes between the mountains. Throughout this region, oak and willow forests primarily border the rivers that flow down from the mountains to the west, and thus dominate the riparian zones, and many of these rivers still flow year-round at mid to high elevations. Overall, the region 1s relatively cool for most of the year, due to the cold California Current that drives winds blowing off the Pacific Ocean waters, bathing the low elevations in morning fog (Hastings and Turner 1965; Markham 1972; Meigs Amphib. Reptile Conserv. 61 Fig. 3. Coniferous forest in Sierra San Pedro Martir, Baja California. Picacho del Diablo (left), the highest peak on the peninsula 1966). Temperatures during the summer months average 20 to 25 °C, whereas winter temperatures average 10.0 to 12.5 °C. The precipitation comes from the tail end of Pacific winter storms that originate from the north, with a prominent decline in rain farther south (Humphrey 1974). This region is a southern extension of the coastal sage scrub and chaparral communities of southern California, which are dominated by sages (Artemisia and Salvia), buckwheat (Eriogonum), lilacs (Ceanothus), and chamise (Adenostoma). Coniferous Forest Region. Prominent forests occur in both the Sierras Juarez and San Pedro Martir (Fig. 3), which extend from the United States in the north and represent the southernmost segment of the Sierran November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula OEE a ike ide Fig. 4. Vegetation in the Lower Colorado Region at Valle del Borrego, Sierra San Felipe, Baja California. Photo by Jorge H. Valdez- Villavicencio. Montane Conifer Forest (Pase 1982). These Peninsular Range mountains are confined to the northern state of Baja California and extend southward from the United States-Mexico border for approximately 300 km. Oriented in a north-south direction, they gradually rise along their western slopes, crest with peaks as high as 3,100 m, and rapidly descend to the desert floor with steep eastern escarpments. The mean monthly air temperatures range from -0.2 to 17 °C. This region has the most reliable precipitation on the peninsula, and receives more rainfall than any other area in Baja California (Hastings and Turner 1965). The majority of the precipitation falls during winter and spring in the form of snow at the higher elevations. Summer brings more variable thunderstorms, when the tropical air moving in a northwesterly direction is uplifted and cooled along the steep eastern slopes of the mountains (Humphrey 1974). The mountains contain extensive meadows that fill with water. The floristic composition of the region is relatively diverse, defined mainly by large shrubs and trees, which are dominated by Jeffrey Pines (Pinus Jeffreyi), manzanitas (Arctostaphylos), oaks (Quercus), firs (Abies), aspens (Populus), and cedars (Calocedrus) (Garcillan et al. 2012). Lower Colorado Desert Region. The Lower Colorado Desert (Fig. 4) is the largest subregion of the Sonoran Desert and extends across southeastern California, southwestern Arizona, northwestern Sonora, and Amphib. Reptile Conserv. 62 northeastern Baja California (Shreve 1951). On the peninsula, this low-elevation desert extends from the Peninsular Ranges to the west and the Colorado River to the east. Confined to the northern state of Baja California, it extends for 450 km southward from the border of the United States to the vicinity of Bahia de los Angeles, where it intergrades widely with the Gulf Coast Desert Region (Peinado et al. 1994). This region contains the late Miocene to Pliocene sedimentary deposits of the Colorado River and the receding waters of the Gulf of California, and is composed of broad expansive basins with elevations ranging from below sea level to 400 m (Spencer and Pearthree 2001). Along this region’s northeastern boundary, the Rio Colorado once provided a rich aquatic ecosystem in the middle of a harsh desert. Today, water no longer flows from the river to the Gulf of California due to its diversion for agricultural and urban use. The mean temperatures for July and August are above 32.5 °C, while the mean winter temperature is 12.5 °C (Markham 1972). Lying in the rainshadow of the Sierras Juarez and San Pedro Martir, this region is the hottest and most barren desert on the peninsula, as it receives less than 5 cm of annual rainfall. Creosote Bush (Larrea tridentata), Burro Weed (Ambrosia dumosa), and Ocotillos (Fouqueria splendens) dominate the vegetation of this region, but it also contains other arid-adapted plants such as mesquites, agaves, palo verde, and various forms of cacti (Gonzalez-Abraham et al. 2010). November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Fig. 5. A view of the Vizcaino Region at Sierra La Asamblea, Baja California, as seen from Transpeninsular Highway 1. Photo by Jorge H. Valdez-Villavicencio. Fig. 6. South of Vizcaino, Baja California Sur; fog is an essential component of this region. Photo by Jorge H. Valdez-Villavicencio. Vizcaino Desert Region. This region (Figs. 5-6) is located in the central portion of the peninsula, and extends from the California and Lower Colorado Desert Regions to Laguna San Ignacio. It 1s bordered by the Pacific Ocean to the west, but its eastern extent varies greatly and generally is limited by the Central Gulf Coast Desert Region that borders the Gulf of California. The southern portion of the Vizcaino Desert Region is much flatter, with elevations just above sea level, whereas the northern portion consists of smaller Amphib. Reptile Conserv. mountain ranges, mesas, and dry washes that are no higher than 1,000 m (Bostic 1971). This region spans both Baja California and Baja California Sur. Many spring-fed oases are found throughout this region and support more mesic communities (Grismer and McGuire 1993). It also experiences a “fog type” temperate desert climate, with limited winter and summer precipitation (Meigs 1966). The mild climate is greatly influenced by prevailing westerly winds coming off the Pacific Ocean, which generate the conditions for heavy fog (Bostic November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula oe Fig. 8. Vegetation in the Villavicencio. 1971). Rainfall occurs in the winter and averages only 5.5 cm. The mean air temperatures range between 23 and 28 °C during the summer, and 15° and 18 °C in winter (Markham 1972). Overall, this region often has overcast skies and a mild climate that receives little rain. In much of the region, the vegetation is open, stunted, widely spaced, and depauperate, because of continuous onshore winds from the Pacific Ocean (Grismer 2002). In areas protected from the winds, plant diversity increases sharply and the dominant plants include Cirios (Fouquieria columnaris), Baja California Tree Yucca (Yucca valida), Cardon (Pachycereus pringlei), Elephant Trees (Pachycormus discolor), mesquites (Prosopis), and agaves (Agave) (Garcillan et al. 2012). Central Gulf Coast Desert Region. This long and narrow region (Figs. 7-9) lies along the eastern coast Amphib. Reptile Conserv. a Central Gulf Coast Region near Bahia Concepcion, Baja California Sur. Photo by Jorge H. Valdez- of the peninsula, extends southward from Bahia de los Angeles to the Cape Region, and spans both states on the peninsula (Shreve and Wiggins 1964). Broad intermixing occurs between this region and the Lower Colorado Desert to the north (Peinado et al. 1994), whereas its boundaries to the west are marked by the uplift of the Peninsular Ranges. The elevation ranges from sea level to 800 m. This region 1s hot and arid, and receives nearly all of its precipitation during the summer and fall. Severe droughts occur in spring, a time when the mean precipitation is only 0.20 cm (Hastings and Turner 1965; Humphrey 1974). The majority of the Central Gulf Coast Desert rainfall originates from southern convectional storms, and appears as run-off from the bordering Peninsular Range. Occasionally, the region receives rainfall from hurricanes that originate November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. a Fig. 10. View of the Magdalena Region at La Purisima, Baja California Sur. Photo by Gerardo Marron. in the southern tropical waters and track northward. Despite periods of severe drought, the annual mean rainfall can reach 16.8 cm (Hastings and Turner 1965) when there has been an active hurricane season. The mean temperatures for the warmest summer months are above 30 °C, whereas the mean winter temperatures fall to around 15 °C (Markham 1972). Arid-adapted plants that include Elephant Trees (P. discolor), Palo Blanco (Lysiloma candidum), \omboy (Jatropha), Palo Adan (Fouquieria diguettii), and various forms of cacti, including large columnar forms, characterize the vegetation. Some mangroves are also present in the southern part of this region, where Black Mangrove (Avicennia germinans) and White Mangrove (Laguncularia racemosa) are the most predominant species (Gonzalez-Zamorano et al. 2011; Garcillan et al. 2012). Amphib. Reptile Conserv. San Basilio, 65 Baja California Sur. Photo : Ss Magdalena Region. This region (Figs. 10-12) extends from Laguna San Ignacio to the Cape Region, along the southwestern coast of the peninsula, and receives the Pacific drainages of the Sierra Guadalupe and Sierra La Giganta. Its eastern border is defined by the uplift of the Peninsular Ranges and its contact with the Central Gulf Coast Desert. This region is confined to Baja California Sur, and is composed of a rugged mountainous region along the eastern border and a flat open plain along the western portion. The mountains contain volcanic badlands interspersed with spring-fed oases (Grismer and McGuire 1993), whereas the western plains consist of sandy fluvial deposits that rely on run-off from the mountains. The elevation in this region ranges from sea level to 1,100 m. The coastal areas receive cool morning fog, but precipitation is low and unpredictable. The mean annual rainfall can reach 12.5 cm (Hastings and Turner November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula " ae tn Fig. 11. View of the Madgalena Region at Los Burros, Baja California Sur. Photo by Bradford Hollingsworth. a a ag aan A Fig. 12. View of the Magdalena Region at 3.4 km NW Puerto Cancun, B 1965). The mean temperatures for the warmest summer months are above 29 °C, whereas the mean winter temperatures are only 17 °C (Markham 1972). Creosote Bush (Larrea tridentata), Elephant Trees (Bursera), Peninsular Palo Verde (Parkinsonia florida), and columnar and various other types of cacti, mesquites, and palms (Washingtonia) in oases dominate the vegetation (Garcillan et al. 2012). Sierra La Laguna Region. The Sierra La Laguna Region (Fig. 13) is located at the southern tip of the Baja California Peninsula, within the Cape Region, and contains dense forests at the higher elevations. Formed by granitic and intrusive rock, these mountains rise to an Amphib. Reptile Conserv. 66 Pe aja California Sur. Photo by Gerardo Marron. elevation of 2,200 m, and have undergone a long history of isolation (Garcillan et al. 2012). Confined to the state of Baja California Sur, this region extends from above 800 m and receives its moisture from summer convectional storms and passing hurricanes. The mean annual rainfall can reach 29.6 cm (Hastings and Turner 1965). Due to a wide range in elevation, the temperatures vary greatly. At the higher elevations, the mean temperatures for the warmest summer months are above 18 °C, whereas the mean winter temperatures are only 8 °C (Markham 1972). The vegetation contains numerous endemics, and the mid-elevation slopes are covered with oak woodlands (Quercus tuberculata,; O. brandegeei), while the upper November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Fig. 13. A view of Sierra La Laguna, the highest mountain range in Baja California Sur, as seen from Segundo Valle. Photo by Jorge H. Valdez-Villavicencio. if Fig. 14. Arid Tropical Region vegetation is evident along Transpeninsular Highway 1, south of La Paz between San Antonio and San Bartolo, Baja California Sur. Photo by Jorge H. Valdez-Villavicencio. elevations are covered with pine-oak woodlands (Q. devia and Pinus lagunae), Peninsular Madrone (Arbutus peninsularis), and Belding Bear-grass (Nolina beldingii) (Garcillan et al. 2012). Arid Tropical Region. This region (Fig. 14) is comprised of the Sierra La Giganta, located along the central uplift of the southern peninsula and the lowlands of the Cape Region, and extends from south of the Isthmus of La Paz to the southern terminus of the peninsula. In the Cape Region, two well-differentiated landscapes of mountain foothills and coastal alluvial plains characterize this region (Garcillan et al. 2012). The foothills extend from 500 to 1,000 m in elevation and intermix with the Sierra La Laguna Region, whereas the alluvial plains are Amphib. Reptile Conserv. 67 found from sea level to 500 m. This region is confined to the state of Baja California Sur. The mean annual rainfall can reach 29.2 cm (Hastings and Turner 1965), and usually occurs in late summer and early fall. This region is hot, with mean temperatures for the warmest summer months above 28.8 °C (Markham 1972). The vegetation is composed of tropical dry forests in the foothills that remain leafless in the dry season, but rebound with the late summer rains. The vegetation includes a variety of woody trees like Palo Blanco (Lysiloma candidum), Mauto (L. divaricatum), Plumeria (Plumeria rubra), and Cardon Barbon (Pachycereus pecten-aboriginum). The coastal lowlands are composed of a fleshy-stemmed shrubland that includes elephant trees (Bursera), Ashy November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula ee au Se, Fig. 15. A view from the southern end of the Todos Santos Sur island in which the northern part of the south island and the Todos Santos Norte island, including its lighthouse, are visible. Photo by Jorge H. Valdez-Villavicencio. Limberbush (Jatropha cinerea), figs (Ficus brandegeei), and succulents (Garcillan et al. 2012). The Pacific Islands. Along the west coast of the peninsula, the Pacific Ocean 1s dominated by the California Current System, which extends from Alaska to northern Baja California, but seasonally extends southward to the tip of the peninsula (Hickey 1979; Badan-Dangon et al. 1989). The California Current consists of a year-round equatorward surface flow that brings cool water down the coastal waters of the peninsula (Kurczyn et al. 2019). On the contrary, the coast of mainland Mexico to the south is exposed to warmer waters from the Equatorial Amphib. Reptile Conserv. Fig. 16. Islas los Coronado (four islands), Baja California. View from Coronado Sur, of Coronado Norte, Pilon de Azucar, and Coronado Medio. Photo by Jorge H. Valdez-Villavicencio. Countercurrent that feeds the Costa Rica Coastal Current flowing poleward, bringing warm counterflows northward along the peninsula (Badan-Dangon et al. 1989). During El Nifio years, the poleward surface currents intensify and bring warmer water farther north, along with some marine reptiles (Grismer 2002). The west coast of the peninsula also contains extensive lagoons and bays which provide important warm-water shelters that are preferred by some sea turtles (Senko et al. 2010). The Pacific Islands (Figs. 15-17) extend along much of the length of the peninsula, from the Islas Coronado offshore from Tijuana, which are located just south of the November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Fig. 17. Todos Santos Sur Island is almost entirely invaded by different species of dense grasses (e.g., Bromus and Avena) and ice plants, which displace the native vegetation. The invasion of introduced grasses and ice plants (Vesembryanthemum crystallinum) occurs throughout the peninsula. Photo by Jorge H. Valdez-Villavicencio. border between California in the United States and Baja California in Mexico, to Isla Creciente, the southernmost island of the insular group, which lies off the Magdalena region of the southwestern peninsula. As noted by Grismer (2002: 9), “the islands of the Pacific coast of Baja California are all landbridge in origin, except for the Islas San Benito, which are oceanic...The largest and most environmentally diverse Pacific island is Isla Cedros, which reaches nearly 1,200 m in elevation. The remaining islands are generally low, small, and rocky, with the notable exception of Isla Creciente, the long, narrow sandbar enclosing the southern end of Bahia Magdalena.” The Gulf Islands. The Gulf of California is a 1,400 km long semi-enclosed sea that extends from a depth of more than 3,000 m at the southern entrance to only Amphib. Reptile Conserv. 200 m at its enclosed end at the Colorado River outlet (Alvarez-Borrego 2002; Lavin and Marinone 2003). The entrance to the Gulf of California is exposed to warmer waters from the poleward flowing Costa Rica Coastal Current (Badan-Dangon et al. 1989). The current in the sea flows northward along the mainland coast and southward down the peninsula during the summer months, and then reverses direction during the winter (Alvarez-Borrego 2002). The mean temperatures in the northern portion of the sea are 8.2 °C in December and 32.6 °C in August (Alvarez-Borrego 2002). Numerous islands and bays (Figs. 18-23) provide a diversity of coastal microhabitats that are favorable to marine reptiles, including sandy beaches, mangrove forests, reefs, and shelters, whereas deep water pelagic conditions exist in the southern Gulf of California, November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula al ee | Yer @ %& a : ai Se ya 3 , cy ee ao > a? Fig. 18. View of the shore of Isla Carmen located in the southern Gulf of California and protected within Parque Nacional Bahia de Loreto. This island is part of the Bahia de Loreto National Park. Photo by Bradford D. Hollingsworth. tees ; Sa ee Fig. 19. View of Puerto Balandra, Isla Carmen, a small bay on the northwestern side of the island, representing the typical arid habitat found within the Gulf Island Region, and home to the single-region endemics Sauromalus slevini and Aspidoscelis carmenensis. Photo by Bradford D. Hollingsworth. which support open-water species (Santamaria-del- in the Baja California Peninsula are given in Table 1. Angel et al. 1994; Lavin and Marinone 2003). The elevations for these localities range from 3 m at Isla Cedros in the Pacific Islands to 1,580 m at Laguna Climate Hanson in the Baja California Coniferous Forest. The mean annual temperature (MAT) 1s highest at Temperature. The monthly minimum, mean, and Loreto (elevation 20 m) in the Central Gulf Coast Region maximum temperatures for one representative locality | and lowest at Laguna Hanson (elevation 1,580 m) in the from each of the 10 geographic regions we recognize — Baja California Coniferous Forest Region. Amphib. Reptile Conserv. 70 November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Fig. 20. View of the uninhabited Isla San Francisco, a small island located in the southern Gulf of California and home to the single- region endemics Aspidoscelis celeripes and A. franciscensis. Photo by Bradford D. Hollingsworth. . a ie a Fig. 21. View of Isla Carmen, Gulf of California, protected within Parque Nacional Bahia de Loreto. Photo by Bradford D. Hollingsworth. Fig. 22. View of the uninhabited Isla Santa Catalina, also known as Catalana Island, located in the Gulf of California off the coast of Loreto, Baja California Sur, which is protected within Parque Nacional Bahia de Loreto and home to some of the most vulnerable species, including nine single-region endemics. Photo by Bradford D. Hollingsworth. Amphib. Reptile Conserv. TA November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Fig. 23. Aerial view of the uninhabited Isla Danzante, which is protected within Parque Nacional Bahia de Loreto, with cliffs up to 150 m, representing the rugged topography typical of islands in the Gulf Island Region. Photo by Bradford D. Hollingsworth. Among the 10 geographic regions of the Baja California Peninsula, the minimum annual temperatures range from 2.2 °C at Laguna Hanson in the Baja California Coniferous Forest to 17.8 °C in three localities: Santo Tomas (180 m) in the California Region, Loreto (20 m) in the Central Gulf Coast Region, and El Barril (50 m) in the Gulf Islands Region. The minimum monthly temperatures peak in either July (Baja California Coniferous Forest Region), August (California Region, Lower Colorado Valley Region, Vizcaino Region, Central Gulf Coast Region, Magdalena Region, Arid Tropical Region, Sierra La Laguna Region, and Gulf Islands Region), or August and September (Pacific Islands Region). The minimum monthly temperatures reach their lowest levels in either January (California, Lower Colorado Valley, Baja California Coniferous Forest, Central Gulf Coast, Magdalena, Arid Tropical, Sierra La Laguna, Pacific Islands, and Gulf Islands regions) or December and January (Vizcaino region). The maximum monthly temperatures are highest in either July (Lower Colorado Valley and Baja California Coniferous Forest regions), August (California, Vizcaino, Central Gulf Coast, Magdalena, Arid Tropical, Sierra La Laguna, and Gulf Islands regions), or September (Pacific Islands region), and are lowest in either January (California, Lower Colorado Valley, Baja California Coniferous Forest, Central Gulf Coast, Magdalena, Arid Tropical, Sierra La Laguna, Pacific Islands, and Gulf Islands regions) or December and January (Vizcaino region). Precipitation. The patterns of precipitation in the Baja California Peninsula are peculiar compared to those Amphib. Reptile Conserv. 72 of other regions in Mexico. Whereas the rainy and dry seasons are confined to specific sets of six months during the year in most areas of Mexico, the rainy season in the Baja California Peninsula is comprised of no more than three or four months. In addition, the months involved are not the same throughout the Peninsula, and are not necessarily sequential. Table 2 provides the precipitation data for 10 localities within each of the 10 geographic regions that we recognize in the peninsula, including the Pacific and Gulf islands. The rainy season extends for three months in the Lower Colorado Valley, the Baja California Coniferous Forest, the Central Gulf Coast, the Magdalena Region, and the Pacific Islands; while it extends for four months in the other five regions. The months involved in the 3-month regions are August through October (in the Lower Colorado Valley, the Central Gulf Coast, and the Magdalena Region), January through March (in the Baja California Coniferous Forest), and December through February (in the Pacific Islands). Those included in the 4-month regions are December through March (in the California Region), September through December (in the Vizcaino Region), August through November (in the Arid Tropical Region), July through October (in the Sierra La Laguna Region), and August through October plus December (in the Gulf Islands). The annual precipitation ranges from 52.2 mm in the Pacific Islands to 581.8 mm in the Sierra La Laguna Region. The mean annual precipitation for the 10 regions is 211.3 mm. The six annual precipitation values lying below this mean are for the Lower Colorado Valley (60.8 mm), the Vizcaino Region (86.0 mm), the Central Gulf Coast Region (160.0 mm), the Magdalena Region (127.0 mm), the Pacific Islands (52.2 mm), and the Gulf Islands November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Table 1. Monthly minimum, mean (in parentheses), maximum, and annual temperature data (in °C) for the 10 geographic regions of the Baja California Peninsula, Mexico. Localities in each region and their elevations are as follows: California, Santo Tomas (180 m); Lower Colorado Valley, El Mayor (15 m); Baja California Coniferous Forest, Laguna Hanson (1,580 m); Vizcaino, Benito Juarez (55 m); Central Gulf Coast, Loreto (20 m); Magdalena, La Purisima (95 m); Arid Tropical, San José del Cabo (20 m); Sierra La Laguna, San Vicente de la Sierra (650 m); Pacific Islands, Isla Cedros (3 m); and Gulf Islands, El Barril (SO m). Data were taken from Servicio Meteorologico Nacional at https://smn.conagua.gob.mx (accessed 18 July 2022). 3 5.7 7.2 9.5 5.6 3.3 n 5 4. 9.5 11.2 13.6 14.3 13.3 California (12.2) (18.3) 49 Lower Colorado Valley (12.6) 20.2 Annual 17.8 (17.8) 27.2 (16.0) 24.7 (14.0) 223 (13.0) 21.3 (20.9) 30.6 (24.1) 34.6 (24.6) 34.8 (23.6) 34.0 (19.5) 29.5 (15.2) 24.8 6 (12.4) 20.9 yep 21.5 6.6 8.5 (14.1) | (16.3) 21.6 | 242 10.4 13 16.1 19.3 (18.7) | (23.8) } (28.6) | (31.3) 27 34.6 43.4 19.8 17.5 13.2 8.8 5.2 11.9 (31.3) | (28.6) | (23) | (17.1) | 2.8) | @1.5) 429 | 396 | 329 | 255 | 204 31.1 9.1 71 Db: -0.8 2.4 ae) (18.5) | (16.2) | (41a) | (74) | (6.4) (10.7) 279 | 254 20 15.7 13.4 19.1 17.0 13.6 9.5 6.9 11.6 (22.9) | (19.5) | (16.7) | (20.9) 32.2 | 294 | 264 30.2 (25.9) 34.9 20.7 15.8 12.2 17.8 (26.9) | (22.0) | (18.3) | (24.0) 33.1 283 | 244 30.2 4] oa 5.6 9.5 (11.5) | (15.1) | (19.0) 20.3 | 246 | 284 Baja Californi -2.7 -2.3 -1.6 0 Coniferous Forest e>) G4) OP (8.0) 11.7 13.1 12.9 16 6.4 7.3 8.8 Vizcaino (16.4) | (17.2) | (18.3) 26.4 27.1 27.9 11.0 11.2 12.4 Central Gulf Coast (17.2) | (17.9) | (19.3) 23.5 24.6 26.3 10.2 (19.5) 28.8 12.0 13.5 16.1 17.5 (20.6) | (22.1) | (25.1) | (26.6) 29.2 | 30.7 | 342 | 357 14.6 (21.8) 28.9 17.6 (24.7) 31.8 21.8 (28.2) 34.6 25.6 (30.7) 35.8 26.0 (31.1) 36.1 24.8 (30.2) 35.5 10.6 12.3 (20.7) | (22.0) 30.7 | 31.8 152 | 20.1 (24.7) | (28.6) 34.2 | 372 21.5 (29.6) 37.7 20.8 (28.9) 37.1 15.9 11.7 9.5 (25.2) | (21.2) } (18.4) 34.4 | 30.7 | 273 13.8 (22.9) 32.1 8.6 9.0 98 Magdalena (17.7) | (18.5) | (19.5) 26.8 28.0 29.2 12.0 12.0 ee (18.9) | (19.2) | (20.1) 258 | 264 | 27.4 6.4 7.2 8.4 Sierra La Laguna (15.7) | (16.5) | (17.7) 24.9 25.8 27.1 143 14.7 14.7 Pacific Islands (18.6) | (19.1) | (19.2) 22.8 10.8 11.5 12.8 Gulf Islands 23.4 | 23.6 (16.4) | (17.3) | (18.8) 219 | 23.1 | 247 (91.6 mm). The four values lying above this mean are for the California Region (274.9 mm), the Baja California 13.4 (20.2) 26.9 17.4 (23.8) 30.2 14.8 17.1 (21.9) | (24.0) 29.1 30.9 20.5 (26.5) 32.6 23.3 (28.5) 33.8 23.8 (28.9) 33.9 23.0 (28.2) 33.4 18.1 (26.2) 34.4 20.5 19.1 17.0 15.1 17.1 (25.0) | (23.8) | (21.4) | (19.2) | (21.5) 295. || =28.6>|| 3257 | 4230 26.0 20.1 16.3 (26.2) | (23.2) 32.3 | 301 Arid Tropical 19.8 (27.9) 36.6 16.0 11.7 8.2 (24.6) | (20.7) | (17.4) 33.2 | 297 | 266 13.0 (22.1) 31.1 10.9 (21.0) 31.1 13.9 16.5 19.4 (23.6) | (25.8) | (27.9) 33.3 | 352 | 364 15.7 16.3 17.3 19.4 (20.5) | (21.0) } (21.7) | (24.0) 25.3 | 257 | 262 | 286 20.5 (24.9) 29.4 20.1 14.8 11.4 (25.8) | (20.6) | (17.0) 31.4 | 264 | 226 17.8 (23.3) 28.8 15.0 (21.0) 27.1 17.7 (23.6) 29.5 22.0 (27.7) 33.4 26.0 (30.7) 35.4 26.3 (30.8) 35.4 24.8 (29.8) 34.7 The range of percentages for the 3-month rainy seasons is 45.1-69.9% while it is 55.6-89.4% for the 4-month Coniferous Forest (390.9 mm), the Arid Tropical Region (288.0 mm), and the Sierra La Laguna Region (581.8 mm). The percentages of annual precipitation that occur during the rainy season are as follows: ¢ California Region: 210.8/274.9 = 76.7% (4-month rainy season) ¢ Lower California Valley: 27.4/60.8 = (3-month) ¢ Baja California Coniferous Forest: 176.4/390.9 = 45.1% (3-month) ¢ Vizcaino Region: 47.8/86.0 = 55.6% (4-month) ¢ Central Gulf Coast Region: 111.9/160.0 = 69.9% (3-month) ¢ Magdalena Region: 66.1/127.0 = 52.0% (3-month) 45.1% rainy seasons. Composition of the Herpetofauna Families The members of the native and non-native herpetofauna of the Baja California Peninsula and its adjacent islands (hereinafter “the Baja California Peninsula” or simply “the peninsula”) are assigned to 32 families, including five families of anurans, one of salamanders, 20 of squamates, and six of turtles (Table 3). The total comprises 51.6% of the 62 families of native and non- native species found in Mexico. No crocodylian or caecilian families are represented on the peninsula. Of the six amphibian families represented on the peninsula, ¢ Arid Tropical Region: 209.0/288.0 = 72.6% 85.7% of the 21 species (Tables 4 and 5) are placed in the (4-month) families Bufonidae (six species), Hylidae (three), Ranidae ¢ Sierra La Laguna Region: 519.9/581.8 = 89.4% (six), and Plethodontidae (three). Among the 26 reptilian (4-month) families, 122 of the 151 species (80.8%) are allocated * Pacific Islands: 30.0/52.2 =57.5% (3-month) in the families Crotaphytidae (five species), Iguanidae ¢ Gulf Islands: 67.4/91.6 = 73.6% (4-month) (nine), Phrynosomatidae (30), Phyllodactylidae (five), Amphib. Reptile Conserv. 73 November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Teiidae (13), Xantusiidae (four), Colubridae (29), Dipsadidae (seven), Natricidae (four), Viperidae (12), and Cheloniidae (four). Page Genera WB Representatives of 64 herpetofaunal genera have been recorded from the Baja California Peninsula, including nine genera of anurans, three of caudates, 42 of squamates, and 10 of turtles (Table 3). These 64 taxa comprise 30.0% of the 213 genera that occur in Mexico (J. Johnson, unpub. data, 27 May 2023). Among the amphibians, the largest numbers of species in Baja California are in the genera Anaxyrus (five species) and Lithobates (four); and among reptiles the largest numbers are in the genera Elgaria (five), Sauromalus (five), Petrosaurus (four), Phrynosoma (five), Sceloporus (10), Urosaurus (four), Uta (five), Phyllodactylus (five), Aspidoscelis (13), Xantusia (four), Lampropeltis (four), Masticophis (five), Sonora (eight), Hypsiglena (six), Thamnophis (four), and Crotalus (12). O59 Species 18.8 The herpetofauna of the Baja California Peninsula consists of 172 species, including 18 anurans, three salamanders, 140 squamates, and 11 turtles (Table 3). The current numbers of native species in these four groups in Mexico are 272, 161, 913, and 51, respectively (J. Johnson, unpub. data, 27 May 2023). The 172 herpetofaunal species known from the Baja California Peninsula represent 12.3% of the 1,397 species of anurans, salamanders, squamates, and turtles in the entire country (J. Johnson, unpub. data, 27 May 2023). The only Mexican state that shares a border with the peninsula is Sonora, and that border is limited in extent. Rorabaugh and Lemos Espinal (2016) reported the number of herpetofaunal species in Sonora as 196, which is 1.1 times the size of the herpetofauna of the peninsula. This proportion is similar to the relative areas of the two regions. The surface area of Sonora is 185,430 km? (Rorabaugh and Lemos Espinal 2016) and that of the Baja California Peninsula is approximately 143,400 km?; thus, Sonora 1s about 1.3 times the size of the entire peninsula. Therefore, the area/species richness ratios are 828.9 for the peninsula and 946.1 for Sonora. Fourteen non-native species comprise 8.1% of total herpetofauna (172 species) of the Baja California Peninsula, and each of the two states contains nine non- native species and share four species. This is the highest number so far in our series of Mexican state herpetofaunas, with an average of four non-natives. Of the 14 non-native species in the Peninsula, five are amphibians, and nine are reptiles (three geckos, two iguanas, one snake, and three turtles), including two species that were listed among the 100 worst invasive alien species (Lithobates catesbeianus and Trachemys scripta) (Lowe et al. 2000). m); and Gulf Islands, El Barril (50 m). Data were taken from Servicio Meteorologico Nacional at https://smn.conagua.gob.mx (accessed 18 July 2022). Shaded areas indicate the months Gulf Coast, Loreto (20 m); Magdalena, La Purisima (95 m); Arid Tropical, San José del Cabo (20 m); Sierra La Laguna, San Vicente de la Sierra (650 m); Pacific Islands, Isla Cedros (3 of the rainy season in each region. Table 2. Monthly and annual precipitation data (in mm) for the 10 geographic regions of the Baja California Peninsula, Mexico. Localities in each region and their elevations are as follows: California, Santo Tomas (180 m); Lower Colorado Valley, El Mayor (15 m); Baja California Coniferous Forest, Laguna Hanson (1,580 m); Vizcaino, Benito Juarez (55 m); Central EN vay [oe S N S | Q S ‘a beara “N + | al Ne) mR Ne) ia) 2 & N — = S oe) Lower Colorado Valley Baja California Coniferous Central Gulf Coast Magdalena Sierra La Laguna Vizcaino Arid Tropical Pacific Islands Amphib. Reptile Conserv. 74 November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Table 3. Composition of the native and non-native herpetofauna of the Baja California Peninsula and its adjacent islands, Mexico. | Order Families Genera Species Anura pe ee eee Cat _) JZ |__| "| Subtotal a 5 |e ee ey] Testudines S____*o,____i Subtotal Sees Patterns of Geographic Distribution We adopted the principal features of the scheme of geographic regions used by Grismer (2002). However, we departed from this system to some extent, in that we recognize the insular regions as separate from the peninsular regions (Table 4). Thus, we recognize eight regions on the peninsula and two regions among the islands (Pacific and Gulf). The distributional data for the 172 species are tabulated in Table 4, and summarized in Table 5 The total number of species in each of the 10 regions (Table 5) ranges from 27 in the Baja California Coniferous Forest Region (BCCFR) to 84 in the Gulf Islands Region (GIR). The average number of regional species is 56.9. The sizes of the herpetofaunas in six of the regions (CR, LCVR, VR, CGCR, ATR, and GIR) are above this average value, and in four (BCCFR, MR, SLLR, and PIR) they are below it. The respective sizes of the regional herpetofaunas do not appear to be related to the relative sizes of the regions themselves, especially since the largest herpetofauna is found on the islands of the Gulf Region. We will examine this issue in greater detail after we present the subsequent analyses. The amphibian fauna of the peninsula 1s comprised of only 21 species (18 anurans and three salamanders), or 12.2% of the total of 172 species. Thus, 87.8% of the total are reptiles. Relatively few of the 151 species of reptiles are turtles, 1.e., 11 species (7.3%). Therefore, most of the herpetofaunal species found in the peninsula are squamates, viz., 140 species, or 81.4% of the total of 172. This result is reasonable, given that the general climate of the peninsula is arid and that much of the diversity of the herpetofauna is insular in distribution (see below). Of the 140 squamate species, 83 (59.3%) are lizards and 57 (40.7%) are snakes. Consequently, the peninsula is a hotspot for lizards, since the 83 species comprise 48.3% of the total herpetofauna. The proportion of the total herpetofauna of 172 species found in each of the 10 regions ranges from 15.9% (27 species in the BCCFR) to 48.8% (84 in the GIR), which indicates the limited extent of species distributions on the peninsula. The members of the peninsular herpetofauna occur in from one to 10 regions as follows: one region (78 of 172 species, 45.3%); two regions (16, 9.3%); three (17, 9.9%); four (10, 5.8%); five (13, 7.6%); six (seven, 4.1%); seven (six, 3.5%); eight (12, 7.0%); nine (7, Amphib. Reptile Conserv. 4.1%), and 10 (six, 3.5%). The mean regional occupancy is 3.3. This figure lies within the range of 1.6 to 3.7 for the other states dealt with thus far in the MCS (Alvarado- Diaz et al. 2013; Mata-Silva et al. 2015: Johnson et al. 2015; Teran-Juarez et al. 2016; Woolrich-Pifia et al. 2016; Nevarez-de los Reyes et al. 2016; Cruz-Saenz et al. 2017; Gonzalez-Sanchez et al. 2017; Lazcano et al. 2019; Ramirez-Bautista et al. 2020; Torres-Hernandez et al. 2021; Cruz-Elizalde et al. 2022). Of the 172 species known from the peninsula, a relatively large proportion (45.3%) is confined to a single region, which is highly significant from a conservation perspective (see below). The number of single-region species ranges from one (in the MR) to 39 (in the GIR). The 39 single-region species in the GIR are as follows, with species endemic to the peninsula indicated by double asterisks, and non-natives indicated by triple asterisks: Crotaphytus insularis** Coleonyx gypsicolus** Dipsosaurus catalinensis** Sauromalus hispidus** Sauromalus klauberi** Sauromalus slevini** Sauromalus varius *** Petrosaurus slevini** Sceloporus angustus** Sceloporus grandaevus** Sceloporus lineatulus** Uta encantadae ** Uta lowei** Uta squamata** Uta tumidarostra** Phyllodactylus bugastrolepis** Phyllodactylus partidus** Aspidoscelis canus** Aspidoscelis carmenensis** Aspidoscelis catalinensis** Aspidoscelis celeripes** Aspidoscelis ceralbensis** Aspidoscelis danheimae** Aspidoscelis espiritensis** Aspidoscelis franciscensis** Aspidoscelis pictus ** Lampropeltis catalinensis** Masticophis barbouri** Rhinocheilus etheridgei** November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 4. Distribution of amphibians and reptiles in the Baja California Peninsula and its adjacent islands, Mexico, by geographic province. Abbreviations: CR = California Region; LCVR = Lower Colorado Valley Region; BCCFR = Baja California Coniferous Forest Region; VR = Vizcaino Region; CGCR = Central Gulf Coast Region; MR = Magdalena Region; ATR = Arid Tropical Region; SLLR = Sierra La Laguna Region; PIR = Pacific Islands Region; and GIR = Gulf Islands Region. * = species endemic to Mexico; ** = species endemic to Baja California; *** = non-native. rr Geographic regions of the Peninsula of Baja California and its adjacent islands Pay a cr [xeve | sccrr | ve | coce | we [are [site [ri | cir | cccupied | Anura(I8species) | TE | Bufonidae(6 species) | | EE cnc ee a A OG GO OO Oe | Anaxyruscalifornicus | + TEE | Anaxyruscognatus TT +! TE | Anaxyrus punctatus | | + OT dT +t UT +h] +h TE +h UT +h +! Ts | Anaxyruswoodhousii_ | | +! TT J Inciliusalvarius TT TP | Hylidae(3speciesy | | Pseudacriscadaverina | + | + | dL TT 8 | Pseudacris hypochondriaca | + | + | + [+] + [+] + [| + | + {[ | 9 | | Smilisca baudinii*** TT | Pipidae(I species) | S| TE Be i a i Cia (en (a (amen a) (mn) (me (un) | Ranidae(6speciesy | | | Lithobates berlandieri*** | | + TE | Lithobates catesbeianus*** | + | + | | + | + TT | Lithobatesforreri*** | | CT CE Ct UT CU +! TCC CC | Lithobatesyavapaiensis | | + | ET | Ranavboyli | Rana draytoni | + [eT | Scaphiopodidae(2speciesy | | | | Scaphiopuscouchii_ | S| + OT dE T+! UT +h] +! UT +! UT CUT +! | Speahammondii | + TT CE TCU | Caudata(3speciesy | TT | Plethodontidae(3species) | | | | Aneidestugubris | + | CUT CUE CC CC | Batrachosepsmajor_ | + | TL 8 | Ensatinaeschscholtzii_—— | + | OT #2 | Squamata(140 species) | | | | Anguidae(Sspecies) | | ET | Elgariacedrosensis** | TT | Elgaria multicarinaa | + | UT TU | Elgarianana**™ | TC CE | Elgariapaucicarinata** || E 8 | Elgaria velazquezi*® | [PP +] +! UT Hh] +! TCU | Anniellidae(2species) | | | | Anniella geronimensis** | + | | dL TT 8 | Anniella stebbinsi_ | +! TT | Bipedidae(I species) | | | Ra 2 en ean Ee eee ee Eee eee Ee ee eee | Crotaphytidae(S species) | | ET | Crotaphytus grismeri** | T+ TP | Crotaphytusinsularis** | TT | Crotaphytus vestigium | + | + | dE HT +! UT +h] +! UT CCU CU ecaribelia copes |i 2 ie ae eee | Gambeliawislizeni | | +! TT | Eublepharidae(3 species) | | | | Coleonyx gypsicolus®* | TE ee a OO | Coleonyxvariegatus | + | + TT + | OT + | + T+ T+ | + | | Gekkonidae(3 species) | | TE | Gehyramutilata*** TT Amphib. Reptile Conserv. 76 November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Table 4 (continued). Distribution of amphibians and reptiles in the Baja California Peninsula and its adjacent islands, Mexico, by geographic province. Abbreviations: CR = California Region; LCVR = Lower Colorado Valley Region; BCCFR = Baja California Coniferous Forest Region; VR = Vizcaino Region; CGCR = Central Gulf Coast Region; MR = Magdalena Region; ATR = Arid Tropical Region; SLLR = Sierra La Laguna Region; PIR = Pacific Islands Region; and GIR = Gulf Islands Region. * = species endemic to Mexico; ** = species endemic to Baja California; *** = non-native. pmo Geographic regions of the Peninsula of Baja California and its adjacent islands Number Taxon of regions |€R | LevR | Becrr | vr | cGcr | MR | ATR | SLLR | PIR | GIR | _ cccupied | Hemidactylusfrenatus*** | | TT Hh THT +] T+ Tt Ts | Hemidactylusturcicus*** | + | + TT | Iguanidae species) | | | Ctenosaurahemilopha** | TT | Dipsosauruscatalinensis** | | | TT | Dipsosaurus dorsalis | | + TT |Iguanarhinolopha*** TT | Sauromalusater PE | Sauromalushispidus** | | | | Sauromalusklauberi** | | Sauromalusstevini®* | Sauromalusvarius®®* PP ea a RS | (ee ee | Callisaurus draconoides | + ere ee a a el Sl | Petrosaurusrepens** PT Ts | Petrosaurus stevini®* |} | TT | Petrosaurus thalassinus** || 8 | Phrynosoma blainvilli | + | | Phrynosomacerroense** | # | TT | Phrynosomacoronatum** | | | TT tT TT +} TU | Phrynosomameallii | | + TT | Phrynosomaplatyrhinos | | + LT | Sceloporusangustus®* PE | Seeloporusgrandaevus®* | | | TT | Sceloporushunsakeri** | TT 8 | Sceloporuslicki** TE | Sceloporuslineatulus*® | | Sceloporusmagister | | + OT TT | Sceloporus occidentalis | + | TT 8 | Sceloporusorcwti tT HT oH Th tT th | Sceloporus vandenburgianus | | TH TT | Sceloporuszosteromus** | + | | dT HT +} UT HT +! T+} Tt Tt 8 | Umanota tt | Urosaurus gracious TE tT | Urosaurustahtelai** TT | Urosaurus nigricaudus | + | + | tT +] +} TP +t] +} T + TP tT +t 8 | Urosaurusornatus TE | Utaencantadae** | stiaiowe tee ee ea eT | Uta squamara®* | Uta tumidarostra*® | Phyllodactylidae(S species) | | | TT | Phyllodactylus bugastrolepis*® | | | TT | Phyllodactylus nocticolus | + | + | tT th | Phyllodactylus partidus*® } | TT | Phyllodactylusunctus** PT | Phyllodactylusxanti*® | | TT TT 8 | Scincidae(3 species) | | | | Plestiodongilberti | + | + Tt UT 8 | Plestiodonlagunensis** | | TT | Plestiodon skiltonianus J] + | | + UT CUT CU a +])+]4+]4 wlelelrmofuofefelelela afeloleluafa Amphib. Reptile Conserv. 77 November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 4 (continued). Distribution of amphibians and reptiles in the Baja California Peninsula and its adjacent islands, Mexico, by geographic province. Abbreviations: CR = California Region; LCVR = Lower Colorado Valley Region; BCCFR = Baja California Coniferous Forest Region; VR = Vizcaino Region; CGCR = Central Gulf Coast Region; MR = Magdalena Region; ATR = Arid Tropical Region; SLLR = Sierra La Laguna Region; PIR = Pacific Islands Region; and GIR = Gulf Islands Region. * = species endemic to Mexico; ** = species endemic to Baja California; *** = non-native. pmo Geographic regions of the Peninsula of Baja California and its adjacent islands Number Taxon of regions | cR | icve | acer | vr | cece | me | arr | site | pm | GIR | _cccupied | Teiidae (13 species) | TT | Aspidosceliscanus** | TT | Aspidoscelis carmenensis** | | | EP | Aspidoscelis catalinensis** | | TE | Aspidoscelisceleripes** | | | EP | Aspidoscelis ceralbensis*® | | | | Aspidoscelis danheimae** | | TE | Aspidoscelis espiritensis*® | | TE | Aspidoscelis franciscensis*® | | TE | Aspidoscelis hyperythrus | + | | TE HT UH! UP +h] + TE + UT +P +] 8 | Aspidoscelistabialis** | * | oT Ee TE | Aspidoscelis maximus** | TT Et TE tT | Aspidoscelis pictus** | | | | CT CT | Aspidoscelistigris J C+ | + {| + | + ] + f+ | ft + | + |g | Xantusiidae (4 species) | | OT | Xanwusiagitberti®® | | Xanwusiahenshawi | *# TT | Xantusia sherbrookei** | TT | Xantusiawigginsi | + | + TCE HT +! TUT CE | Charinidae(Ispeciesy | | | | Lichanuratrivirgata | + | + | | + { + | +] + | + [+] +] 9 | | Colubridae 29 species) | | TP | Arizonaelegans | + | +} OT CET 8 | Arizonapacata** | TT PT | Bogertophisrosaliae | | + OT CPE tT Hh UP th] +} UT + Tt TT | Lampropeltis catalinensis** | | TE | Lampropeltishherrerae** | | | EP | Lampropeltis multifasciaca | | | + | | Masticophis aurigulus** | | TE 8 | Masticophis barbouri** | | | | Masticophis flagellum | | * TE | Masticophis fuliginosus | * | + | + |+ to + P+] + | + f+ >t] | | Masticophistareralis | * | + | + P+ ft + P+} ot ft of ot 6 | | Phyllorhynchus decurtatus | | + TE HT HT +! Te! TU | Pituophis catenifer | + | + [ + | +] | of ot t+*i of 5 | | Pituophis insulanus*® | TT Re rn eS a a ee ae ee ee ee | Rhinocheilus etheridgei** | | TE | Rhinocheilusteconei_ | * | + [ot +f 8 | Salvadorahexalepis | + | + | + P+] + P+} + | + f+? +e | | | Sonoraannuata | | + TE | Sonoracincta** E+ | + TU TH | Sonorafasciata** | | OP | tt Th Pt | Sonoramosaueri®* | | TP eT Hh Ph | +! TU | Sonora punctatissima®® | | TE | Sonorasavagei** PT PE | Sonorastraminea** | TT Et Tt 8 | Sonorasemiannaa | | + OT | Tantillaplaniceps | + | + [| dT +t] +! UP Lh }hU] +} T+ TC UT cv} Ts | Trimorphodon lyrophanes, | + | + | + | +] + J+] +{+ t{ ~P+t »¢ | ee ee ee ee ee eluol—l—Ta>ual[uf— Amphib. Reptile Conserv. 78 November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Table 4 (continued). Distribution of amphibians and reptiles in the Baja California Peninsula and its adjacent islands, Mexico, by geographic province. Abbreviations: CR = California Region; LCVR = Lower Colorado Valley Region; BCCFR = Baja California Coniferous Forest Region; VR = Vizcaino Region; CGCR = Central Gulf Coast Region; MR = Magdalena Region; ATR = Arid Tropical Region; SLLR = Sierra La Laguna Region; PIR = Pacific Islands Region; and GIR = Gulf Islands Region. * = species endemic to Mexico; ** = species endemic to Baja California; *** = non-native. us Geographic regions of the Peninsula of Baja California and its adjacent islands ea a cr | neve | peer | vr | coer | Mm {AR | Stn | PIR} GIR accupied | Dipsadidae (7 species) | | S|) a | CV 2 | Hypsiglenacatalinae** | | TE | Hypsiglenachlorophaea | | + | dE | Hypsighena gularis** | | TE | Hypsighena marcosensis** | | | ET | Hypsighenastevinis® | | + TE + T+ T+ T+! LT +! UT c+] +! Ts | Elapidae(I species) | | | Hydrophisplaturus | | + OT ET + T+! TCU T+ | Leptotyphlopidae(2 species) | | | | ee a) ET (CR (I | Rena humilis f+ OT +) TP Te + TT T+ | Natricidae (4 species) | S| | | Thamnophiselegans | S| TE UT | Thamnophishammondii_ | + | | + oT + T+ UT +h T+! TCU | Thamnophis marcianus | | TL | Thamnophisvalidus** | | TE | Typhlopidae(I species) | | | TT | Indotyphlops braminus*** | + | tT 8 | Viperidae(I2speciesy | | | CT | Crotalus angelensis** | | TE en 5 OO A OO OO | Crotalus catalinensis** | | TE | Crotalus cerastes, | | *# OT E Peretatusizmor IE Weve eile a ee Se ee ee Pe | Crotalushelleri | + | + UT hUv*®hUdT HUT CCU | Crotalus lorenzoensis** | | | CE CT | Crotalus mitcheltii** | | dT EH TH THT +! T+! THT +H 7 | Crotaluspolisi** | | TE CE | Crotalus pyrrhus tT + LT +t UT hth UT HUT CU [sCratalisrubers a ae es Are fete es ee tos | Crotalus thalassoporus** || TE | Testudines(I1 species) | S| | | Cheloniidae (4species) | | | ea cs OO a Oe -Cheloniamdas _ ff PP ers | EO | Dermochelyidae(1 species) | | | | Dermochelyscoriacea | | oO * OT ET TS | Emydidae(3species) | | CT | Actinemys pallida | + | CUT CUE TC | Trachemysnebulosa** | | dT CE ET UT +H T+! UT +! UT CUS | Trachemys scripta*** | + | TE | Kinosternidae (I species) | | | | Kinosternonintegrum*** | | E | Testudinidae(Ispeciesy | | | | Gopherus morafkai_ | | dT | Trionychidae(I species) | | | Sy SEE a) | Apatone spiniferav** | | + OT CE | Total(172speciesy) | | CE Amphib. Reptile Conserv. 79 November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 5. Summary of distributional occurrence of herpetofaunal families in the Baja California Peninsula, Mexico, by geographic region. Abbreviations: CR = California Region; LCVR = Lower Colorado Valley Region; BCCFR = Baja California Coniferous Forest Region; VR = Vizcaino Region; CGCR = Central Gulf Coast Region; MR = Magdalena Region; ATR = Arid Tropical Region; SLLR = Sierra La Laguna Region; PIR = Pacific Islands Region; and GIR = Gulf Islands Region. Number of Distributional occurrence PButonidee | 6 | 2]os | 1 |2) 3: |i | 31 | «1 |i ft, pipe es SE (oe (In see | i Lae I sees —— 0) | |e Oe |e el aa ar a a a FScaphiopodine | 2 | 1 | 1 | — | 2] 1 [|i [| 1 | 1 | —| 11 subtom ——s| te] | | *} ld] 7 | cs | 8 | cs | #2 2 EES AE REE = T= Sipe | > | =m | | | | [ET 7 Jr (RT (al (C(O (ES ( / a ET OS ES aa a | a YE a Pe Fa a a lS (a A ( (comin I Sele | os Look 2 il a) © [= Ls | [Eublepharide | 3 | 1 | 2 | — | 2 | 2 | 1 | 1 | «1 | 1/2 _ Fee SE (O(N | | YS en eee a ee | ee or ee O| oe) oh TlieteN FPhiynosomatae | 30 | 9 | | 4 | 9 | s | « | | s | 6 | 17 FPhyltodacytidae | 5 | 1 | 1 | — | 1 | 3 | 1 | 3 | 2 | 1 | 4 _ semis | | eR | es | io | Le a aa ee a PS aR (a a | fa Ee A AS ee Tsubo «| ep | ow | le | | oe Pl | le) |) ESE] TR OG = (SE CC Fcoubrine | 29 |u| is | 7 | |“ | oe | o | 9 | 9 | «| a Sa ea ee ee vipers | 2 [«| «se | 3 [s5|[ 4 [3 [3 [3 | «| 01] SE LST RC GC Ppermocneyites | 1 |—-|1 | —[-,—-}.1]14]|-—- |]. BA a ST OO T(E ite ie | eee ee | aa oe a a | a as a a (a a eal (| |] See a S05 | a | || | Trot i ise fo | | a [| 7 |» | o | » | o | 2 | Tsumoat | 12 [ol] «| 7 [es] e« [sa | « | «a |« |» | Sonora punctatissima** These species are all peninsular endemics, except for one Sonora savagei** (Sauromalus varius) which was introduced to a small Hypsiglena catalinae** islet included in the Gulf Island group (Hollingsworth et Hypsiglena gularis** al. 1997). These species are all either lizards (26 species) Hypsiglena marcosensis** or snakes (13 species). Crotalus angelensis** The peninsular region with the next largest number of Crotalus catalinensis** single-region species is the LCVR, with 19, as follows Crotalus lorenzoensis** (the number following the species name indicates which Crotalus polisi** of the distributional categories is involved): Crotalus thalassoporus** Amphib. Reptile Conserv. 80 November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Anaxyrus cognatus 3 Anaxyrus woodhousii 3 Incilius alvarius 3 Lithobates berlandieri***2 Lithobates yavapaiensis 3 Crotaphytus grismeri** Gambelia wislizenii 3 Phrynosoma mceallii 3 Phrynosoma platyrhinos 3 Sceloporus magister 3 Uma notata 3 Urosaurus graciosus 3 Urosaurus ornatus 3 Masticophis flagellum 3 Sonora annulata 3 Sonora semiannulata 3 Hypsiglena chlorophaea 3 Thamnophis marcianus 7 Apalone spinifera*** Of these 19 species, 16 (84.2%) are non-endemics, two are introduced species, and one is a peninsular endemic. Of the 16 non-endemic species, all but one are distributed to the north in the United States; and the single exception is the garter snake Yhamnophis marcianus, which occurs from the United States through Mexico, and into Central America (http://mesoamericanherpetology.com; accessed 5 June 2022). Crotaphytus grismeri is endemic to the LCVR, and the two introduced species are from populations outside of Baja California. The third-largest group of single-region species is found in the ATR region, and is comprised of the following five species: Smilisca baudinii*** Gehyra mutilata*** Iguana rhinolopha*** Kinosternon integrum*** Gopherus morafkai 3 Interestingly, four of these five species are introduced, either from elsewhere in Mesoamerica or from outside of Mesoamerica; and the remaining species also 1s distributed in the United States. Three regions contain four single-region species. One of these regions is the CR, and the species are as follows: Anaxyrus californicus 3 Xenopus laevis*** Phrynosoma blainvillii 3 Trachemys scripta*** Two of these species are non-native, and the other two also are distributed in the United States. The second region with four single-region species is the BCCER, and the species are as follows: Amphib. Reptile Conserv. 81 Rana boylii 3 Sceloporus vandenburgianus 3 Lampropeltis multifasciata 3 Thamnophis elegans 3 All four of these species also are distributed to the north in the United States. In significant contrast to the GIR, the PIR supports only four single-region species, as follows: Elgaria cedrosensis** Elgaria nana** Lampropeltis herrerae** Pituophis insulanus** All of these species are peninsular endemics, like most of the species 1n the GIR. Three other regions contain only one single-region species. One is the MR, and the species involved is: Xantusia sherbrookei** This species is a peninsular endemic. The second region with one single-region species is the SLLR, and the species is: Xantusia gilberti** It also is a peninsular endemic. The last region with one single-region species is the VR, and the species is: Urosaurus lahtelai** This species is another peninsular endemic. In summary, of the 78 single-region species found on the peninsula, 46 are peninsular endemics, 23 are non- endemics, and nine are non-native species. Only one of the 10 regions, the CGCR, has no single-region species. Of the 10 phytogeographic regions on the peninsula, the GIR is the most significant with regard to conservation importance, since it contains the largest overall number of species (84), the largest number of single-region species (39), and the greatest number of peninsular endemics (50). Regional Occupancy and the Coefficient of Biogeographic Resemblance (CBR) Another indication of conservation significance involving the 10 phytogeographic regions is the relative average regional occupancy (Table 6). This figure is calculated by recording the number of species occupying each of the regions | through 10. For example, the CR contains 60 species that occupy regions 1—10 as follows: November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 6. Numbers of species, regional occupancy, and average regional occupancy for the geographic regions of the Baja California Peninsula and adjacent islands, Mexico. See Table 4 for abbreviations. ie ee [El [a EE ee PIR GIR CR (60 species) Region | =4 Region 6 = 4 Region 2 = 10 Region 7 = 2 Region 3 = 10 Region 8 = 8 Region 4 =3 Region 9 =7 Region 5 =6 Region 10 = 6 Based on these data, the mean regional occupancy value for the CR is 5.4 (321/60). The mean regional occupancy values for the 10 regions range from 4.1 to 6.9 (Table 6), as follows (in numerical order): GIR = 4.1 VR=6.2 LEVR= 5.0 PIR = 6.3 BCCFR = 5.2 CGCR = 6.4 CR=5.4 SLLR = 6.4 ATR=5.8 MR =6.9 The regional occupancy values roughly indicate the relative conservation significance of each of the 10 regions. Thus, the GIR evidently is the most conservation significant region in the Baja California Peninsula, and the MR is the least. Thus, even though the GIR is the region with the highest herpetofaunal figure (84), it supports the highest degree of single-region species, the peninsular endemic species. As in other MCS studies, we constructed a Coefficient of Biogeographic Resemblance (CBR) matrix in order to elucidate the similarity relationships among the 10 phytogeographic regions we recognize in the Baja Amphib. Reptile Conserv. 82 321 DD, 141 394 California Peninsula, including its associated islands on both the Pacific and Gulf sides (Table 7). The greatest species richness is contained in the Gulf Island Region (84 species), and the least is in the Baja California Coniferous Forest Region (27 species). The number of shared species between each regional pair ranges from eight between the BCCFR (27 species) and SLLR (41 species), which are relatively small areas located roughly at opposite ends of the peninsula and contain relatively small numbers of species, to 54 between the CGCR (62 species) and ATR (65 species), which are relatively low-elevation regions containing relatively high species numbers and are contiguous in the Cape Region of the peninsula. The mean value of shared species among all 10 regions is 29.9. The following data show the ranges and means of shared species (bold in parentheses) for each of the 10 regions, and are arranged according to decreasing species richness (underlined values) in each region: GIR (84): 9-41 (30.3) LCVR (67): 15-43 (30.4) ATR (65): 10-54 (35.4) VR (64): 16-48 (37.1) CGCR (62): 11-54 (37.3) CR (60): 18-45 (29.3) MR (51): 11-48 (34.1) PIR (47): 13-33 (27.4) SLLR (41): 8-39 (24.9) BCCER (27): 8-23 (12.9) The lowest number of shared species in Table 7, 1e., eight between the BCCFR and the SLLR, is understandable inasmuch as six species occur in all 10 of the phytogeographic regions. These six species are Uta stansburiana, Lampropeltis californiae, Masticophis fuliginosus, — Salvadora hexalepis, — Hypsiglena ochrorhynchus, and Crotalus ruber (Table 4). The other two species that occur in nine regions are Pseudacris hypochondriaca and = Trimorphodon _ lyrophanes. November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Table 7. Pair-wise comparison matrix of Coefficient of Biogeographic Resemblance (CBR) data of the herpetofaunal relationships for the 10 physiographic regions in the Baja California Peninsula, Mexico. Underlined values = number of species in each region; upper triangular matrix values = species in common between two regions; and lower triangular matrix values = CBR values. The formula for this algorithm is: CBR = 2C/N, + N, (Duellman 1990), where C is the number of species in common to both regions, N, is the number of species in the first region, and N, is the number of species in the second region. See Table 4 for abbreviations. See Fig. 24 for the UPGMA dendrogram produced from the CBR data. [era [econ [ve [coe | we [| am | se] me [| or | Ce | o« [| « | >» | os |» | «| » | ew | » | 4 Coes [oss | @ [os |» | 7 | =» [| » | » [| 7 |» Pacer [oss | 02 | @ | [|u| u [0 | 3s [| 5 | > vr | o7s [06 | oss | [| # | «= [| © | 2» [| » | 3 [ame [002 [032 | 022 | 06s [oss | oso [os |» | 0 [ a re [om [oar [oss] oss [os | om [oss | oas | a7 [28 [oe [oss [0 [ois | oar [oss [oss [oss | oas | 04s [at Interestingly and perhaps expectedly, only one of the eight is an amphibian, one is a lizard, and the remaining Six are snakes. In addition, the two insular regions positioned on either side of the peninsula (PIR and GIR) might be expected to share relatively few species. Their number of shared species (28) is higher than the number between the BCCFR and the SLLR (Table 7). These 28 species include the six occurring in all 10 regions, as well as one in four regions, two in five regions, one in six regions, three in seven regions, 10 in eight regions, and five in nine regions. Notably, no insular endemic species are shared between these two insular regions. The six peninsular endemic species found in these two insular regions also are found on the intervening mainland regions, and in total are found in five to nine regions. The CBR values in Table 6 range from 0.16 between the BCCFR and the GIR to 0.86 between the ATR and the CGCR. These relationships easily are understood given that the BCCFR is a “cool mesic area” occupying the “upper elevations of the northern Sierra Juarez and southern Sierra San Pedro Martir...” (Grismer, 2002: 12) and the GIR is comprised of the islands in the Gulf region. In addition, the ATR and the CGCR are two regions in the southern portion of the peninsula that are broadly contiguous and overlapping. UPGMA dendrogram The UPGMA dendrogram (Fig. 24) indicates that the two most closely related regions are the adjacent and overlapping Central Gulf Coast Region (CGCR) and the Arid Tropical Region (ATR), which are joined at the 0.86 level. These two regions are joined at the 0.82 level with the Magdalena Region (MR), which is adjacent with both of the Central Gulf Coast and Arid Tropical Regions for some distance. These three regions are joined at the 0.74 level with the Vizcaino Region (VR), which is adjacent Amphib. Reptile Conserv. to both the Central Gulf Coast and Magdalena Regions. These four regions are joined at the 0.64 level with the Sierra La Laguna Region (SLLR), which is located in the southern cape region of Baja California Sur and surrounded by the Arid Tropical Region. This group of five regions in the southern portion of the Peninsula is joined to the remaining regions in the northern portion of the Peninsula and those in the Pacific Ocean and the Gulf of California at the 0.52 level. Of the remaining five regions, the regions most closely allied are the Pacific Islands Region and the California Region, joined at the 0.62 level. These two regions are united to the other three regions at the 0.51 level with the Lower Colorado Valley Region (LCVR). The eight previously mentioned peninsular regions are joined to the Gulf Islands Region at the 0.44 level. Finally, and notably, the most distantly related region is the Baja California Coniferous Forest Region, which 1s joined to all the other regions at the 0.33 level and is “the southernmost disjunct and depauperate section of the broader and more inclusive Sierra Montane Conifer Forest” (Grismer 2002: 11-12). This region has the smallest herpetofauna (27 species) of the 10 regions, and the lowest average number of species in common (12.9) with the remaining regions (see above and Table 7). Distribution Status Categorizations We used the system developed by Alvarado-Diaz et al. (2013) to discuss the distribution status of the members of the Baja California herpetofauna, as was used in the previous studies in the MCS (see above). The categories in this system have been somewhat adapted to include the following: non-endemic, peninsular endemic, and non-native. The categorizations for each species are listed in Table 8, and summarized in Table 9. The species numbers in each of the three distribution categories, in decreasing order, are: non-endemics, 81 (47.1%); peninsular endemics, 77 (44.8%); and non- November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula oa + & & ; ah é a “ 0.9 0.86 0.82 0.8 0.74 0.7 0.64 g 0.62 § q 0.6 wn 0.52 0.51 0.5 0.44 0.4 0.33 Fig. 24. UPGMA generated dendrogram illustrating the similarity relationships of species richness among the herpetofaunal components in the 10 geographic regions of the Baja California Peninsula (based on the data in Table 6; Sokal and Michener 1958). Similarity values were calculated using Duellman’s (1990) Coefficient of Biogeographic Resemblance (CBR). natives, 14 (8.1%). These distribution categories differ from those utilized in other MCS studies, inasmuch as the Baja California Peninsula is almost completely segmented from the remainder of the country of Mexico, and biogeographically it is much more closely associated with the US state of California (Mata-Silva et al., In Press). Therefore, it is not possible to separate country endemics from state endemics, as was done in the other MCS studies, but rather they are recognized here as only a single category, i.e., the peninsular endemic category (Table 7). As expected, almost all of the non-endemic species (74, or 91.4% from a total of 81 species) are categorized as NE3 or MXUS species (1.e., species occurring in both Mexico and the United States), according to the categories established by Wilson et al. (2017). However, one species (1.2%) is an NE7 or USCA species (Thamnophis marcianus), 1.€., a Species occurring from the United States to Central America); and six (7.4%) are NE9 or OCEA species (one marine snake, Hydrophis platurus, and five marine turtles, Caretta caretta, Chelonia mydas, Eretmochelys imbricata, Lepidochelys olivacea, and Dermochelys coriacea). The peninsular endemic species amount to 77 and occupy from one to nine geographic regions (Table 7): one region (46 species, 60.5%); two (four, 5.3%); three (eight, 10.5%); four (five, 6.6%); five (eight, 10.5%); six (none); seven (two, 2.6%); eight (three, 3.9%); and nine (one, 1.3%). The 46 species confined to a single geographic region are either mostly limited to the Pacific Insular Region (four species) or the Gulf Insular Region (38 species). The following four species are confined to the Pacific Insular Region: Amphib. Reptile Conserv. Elgaria cedrosensis** Elgaria nana** Lampropeltis herrerae** Pituophis insulanus** The 38 species limited to the Gulf Insular Region are: 84 Crotaphytus insularis** Coleonyx gypsicolus** Dipsosaurus catalinensis** Sauromalus hispidus** Sauromalus klauberi** Sauromalus slevini** Petrosaurus slevini** Sceloporus angustus** Sceloporus grandaevus** Sceloporus lineatulus** Uta encantadae ** Uta lowei** Uta squamata** Uta tumidarostra** Phyllodactylus bugastrolepis** Phyllodactylus partidus** Aspidoscelis canus** Aspidoscelis carmenensis** Aspidoscelis catalinensis** Aspidoscelis celeripes** Aspidoscelis ceralbensis** Aspidoscelis danheimae** Aspidoscelis espiritensis** Aspidoscelis franciscensis** Aspidoscelis pictus ** November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Table 8. Distributional and conservation status measures for members of the herpetofauna of the Baja California Peninsula, Mexico. Distributional status: PE = endemic to Peninsula of Baja California; NE = not endemic to peninsula; and NN = non-native. The numbers suffixed to the NE category signify the distributional categories developed by Wilson et al. (2017) and implemented in the taxonomic list at the Mesoamerican Herpetology website (http://mesoamericanherpetology.com), as follows: 3 (species distributed only in Mexico and the United States); 4 (species found only in Mexico and Central America); 6 (species ranging from Mexico to South America); 7 (species ranging from the United States to Central America); 8 (species ranging from the United States to South America); and 9 (Oceanic species). Environmental Vulnerability Score (taken from Wilson et al. 2013a,b): low (L) vulnerability species (EVS of 3-9); medium (M) vulnerability species (EVS of 10-13); and high (H) vulnerability species (EVS of 14-19). IUCN categorization: CR = Critically Endangered; EN = Endangered; VU = Vulnerable; NT = Near Threatened; LC = Least Concern; DD = Data Deficient; and NE = Not Evaluated. SEMARNAT status: A = Threatened; P = Endangered; Pr = Special Protection; and NS = No Status. See Alvarado-Diaz et al. (2013), Johnson et al. (2015a), and Mata-Silva et al. (2015) for explanations of the EVS, IUCN, and SEMARNAT rating systems. Taxon Anaxyrus boreas Anaxyrus californicus Anaxyrus cognatus Anaxyrus punctatus Anaxyrus woodhousii Incilius alvarius Pseudacris cadaverina Pseudacris hypochondriaca Smilisca baudinii*** Xenopus laevis*** Lithobates berlandieri*** Lithobates catesbeianus*** Lithobates forreri*** Lithobates yavapaiensis Rana boylii Rana draytonii Scaphiopus couchii Spea hammondii Aneides lugubris Batrachoseps major Ensatina eschscholtzii Elgaria cedrosensis** Elgaria multicarinata Elgaria nana** Elgaria paucicarinata** Elgaria velazquezi** Anniella geronimensis** Anniella stebbinsi Bipes biporus** Crotaphytus grismeri** Crotaphytus insularis** Crotaphytus vestigium Gambelia copeii** Gambelia wislizenii Coleonyx gypsicolus** Coleonyx switaki Coleonyx variegatus Gehyra mutilata*** Hemidactylus frenatus*** Hemidactylus turcicus*** Ctenosaura hemilopha** Dipsosaurus catalinensis** Dipsosaurus dorsalis Amphib. Reptile Conserv. Distributional status i E E E E E E j 3 5) 2 3 3 3 3 3 E E Z Z Z Zlzazilzizlz\lz\|Z E ich 18) E j 3 3 3 3 ES ES NE3 2 E E P P P P N P P P E EB E ie iE E E re E 13) iE N N E E 3 3 3 3 3 NE3 3 FE. E E N NN N P P NE3 Environmental Vulnerability Category (Score) ——— ———— _——— 1 —— | a. | ——<——=—— 85 IUCN categorization LC N Le LG LC LG, LC LG L LC ill Le LG LC C LC IG LC LC N RG, LC LG LC C LC LG It L E c NT NT N L E NE LC 1 C C NE LC status P Pr S S S S S S S S S S S Pr T N T T T N a Se | P. P P P November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 8 (continued). Distributional and conservation status measures for members of the herpetofauna of the Baja California Peninsula, Mexico. Distributional status: PE = endemic to Peninsula of Baja California; NE = not endemic to peninsula; and NN = non-native. The numbers suffixed to the NE category signify the distributional categories developed by Wilson et al. (2017) and implemented in the taxonomic list at the Mesoamerican Herpetology website (http://mesoamericanherpetology.com), as follows: 3 (species distributed only in Mexico and the United States); 4 (species found only in Mexico and Central America); 6 (species ranging from Mexico to South America); 7 (species ranging from the United States to Central America); 8 (species ranging from the United States to South America); and 9 (Oceanic species). Environmental Vulnerability Score (taken from Wilson et al. 2013a,b): low (L) vulnerability species (EVS of 3-9); medium (M) vulnerability species (EVS of 10-13); and high (H) vulnerability species (EVS of 14-19). IUCN categorization: CR = Critically Endangered; EN = Endangered; VU = Vulnerable; NT = Near Threatened; LC = Least Concern; DD = Data Deficient; and NE = Not Evaluated. SEMARNAT status: A = Threatened; P = Endangered; Pr = Special Protection; and NS = No Status. See Alvarado-Diaz et al. (2013), Johnson et al. (2015a), and Mata-Silva et al. (2015) for explanations of the EVS, IUCN, and SEMARNAT rating systems. T Distributional Environmental Vulnerability IUCN SEMARNAT axon at status Category (Score) categorization status Iguana rhinolopha*** ne M (13) C cl [ay Odd a a —— aa a i Sauromalus ater | NEB. Sauromalus hispidus** Ea PES | Sauromalus klauberi** | PECd Sauromalus slevini** | PECd [es [es rey [rey Phrynosona blaimilti | NES Ss M(2)——SS—S —) a _—— | NES | NE ee |e PE een © ay a a —_— | NES | NES Callisaurus draconoides C Petrosaurus mearnsi s Petrosaurus repens** Pr N N N N Phrynosoma cerroense** H (14) M (12) N H (15) N M (13) H (16) H(17) H (14) M (13) H(17) N L(9) Sceloporus occidentalis N M (11) N N N N N N N N Phrynosoma coronatum** Phrynosoma mcallii Phrynosoma platyrhinos Sceloporus angustus** Sceloporus grandaevus** Sceloporus hunsakeri** Sceloporus licki** Sceloporus lineatulus** Sceloporus magister Sceloporus vandenburgianus | NEBL t—“‘é‘aCCAN [rey [es a a or: ss a Uta lowei™ rey | NES La a == | NES LL Eee | a [-_*PEe_s| | NES ed Sceloporus zosteromus** Uma notata Urosaurus graciosus Urosaurus lahtelai** Urosaurus nigricaudus Urosaurus ornatus N Uta stansburiana Uta tumidarostra** Phyllodactylus bugastrolepis** Phyllodactylus nocticolus Phyllodactylus partidus** Phyllodactylus unctus** Phyllodactylus xanti** E V N i N L N L L N V V 1 V N NN E3 PE PE PE NN BS B3 PE PE PE E3 PE PE E3 E3 PE PE PE PE PE YE3 YE3 YE3 PE YE3 YE3 PE YE3 YE3 PE PE PE ES PE PE E3 PE PE PE Plestiodon gilberti ES PE iG, L Le LC LC LC E © T TAS C LC Le LG LC C LC LG LC LG. il LG LC KG, LC U U C Le U LC LG, Le it LC LC LC Plestiodon lagunensis** Amphib. Reptile Conserv. 86 November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Table 8 (continued). Distributional and conservation status measures for members of the herpetofauna of the Baja California Peninsula, Mexico. Distributional status: PE = endemic to Peninsula of Baja California; NE = not endemic to peninsula; and NN = non-native. The numbers suffixed to the NE category signify the distributional categories developed by Wilson et al. (2017) and implemented in the taxonomic list at the Mesoamerican Herpetology website (http://mesoamericanherpetology.com), as follows: 3 (species distributed only in Mexico and the United States); 4 (species found only in Mexico and Central America); 6 (species ranging from Mexico to South America); 7 (species ranging from the United States to Central America); 8 (species ranging from the United States to South America); and 9 (Oceanic species). Environmental Vulnerability Score (taken from Wilson et al. 2013a,b): low (L) vulnerability species (EVS of 3-9); medium (M) vulnerability species (EVS of 10—13); and high (H) vulnerability species (EVS of 14-19). IUCN categorization: CR = Critically Endangered; EN = Endangered; VU = Vulnerable; NT = Near Threatened; LC = Least Concern; DD = Data Deficient; and NE = Not Evaluated. SEMARNAT status: A = Threatened; P = Endangered; Pr = Special Protection; and NS = No Status. See Alvarado-Diaz et al. (2013), Johnson et al. (2015a), and Mata-Silva et al. (2015) for explanations of the EVS, IUCN, and SEMARNAT rating systems. Distributional Environmental Vulnerability IUCN status Category (Score) categorization status N NS P Taxon Plestiodon skiltonianus Aspidoscelis canus** Aspidoscelis carmenensis** Aspidoscelis catalinensis** Aspidoscelis celeripes** Aspidoscelis ceralbensis** Aspidoscelis danheimae** Aspidoscelis espiritensis** Aspidoscelis franciscensis** Aspidoscelis hyperythrus N Aspidoscelis labialis** Aspidoscelis maximus** P P P P P P P P P te P Aspidoscelis pictus** Aspidoscelis tigris P Xantusia gilberti** Xantusia henshawi i Xantusia sherbrookei** Xantusia wigginsi Lichanura trivirgata Arizona elegans Arizona pacata** P Bogertophis rosaliae Lampropeltis californiae Lampropeltis catalinensis** P Lampropeltis herrerae** ly Lampropeltis multifasciata N Masticophis aurigulus** P Pp. Masticophis barbouri** Masticophis flagellum Masticophis fuliginosus Masticophis lateralis Phyllorhynchus decurtatus Pituophis catenifer Le Le LG VU Le LC Le LC LC Le VU NE Le Le NE LC NE NE Le. LC LC IG Le’ Le NE Le. LC NE LC Le. LC Pituophis insulanus** FP LG Pituophis vertebralis** P Ee Rhinocheilus etheridgei** ‘ig Rhinocheilus lecontei Le KG Salvadora hexalepis N N N Sonora annulata N es Sonora cincta** es Sonora fasciata** es P P E3 E E E E E E E E E3 E E E E3 E E3 E E3 E3 E E3 E3 E E E3 E E E3 E3 E3 E3 E3 E E E E3 E3 E3 E3 E Sonora mosaueri** E E P Z ™ Sonora punctatissima** Amphib. Reptile Conserv. 87 November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 8 (continued). Distributional and conservation status measures for members of the herpetofauna of the Baja California Peninsula, Mexico. Distributional status: PE = endemic to Peninsula of Baja California; NE = not endemic to peninsula; and NN = non-native. The numbers suffixed to the NE category signify the distributional categories developed by Wilson et al. (2017) and implemented in the taxonomic list at the Mesoamerican Herpetology website (http://mesoamericanherpetology.com), as follows: 3 (species distributed only in Mexico and the United States); 4 (species found only in Mexico and Central America); 6 (species ranging from Mexico to South America); 7 (species ranging from the United States to Central America); 8 (species ranging from the United States to South America); and 9 (Oceanic species). Environmental Vulnerability Score (taken from Wilson et al. 2013a,b): low (L) vulnerability species (EVS of 3-9); medium (M) vulnerability species (EVS of 10-13); and high (H) vulnerability species (EVS of 14-19). IUCN categorization: CR = Critically Endangered; EN = Endangered; VU = Vulnerable; NT = Near Threatened; LC = Least Concern; DD = Data Deficient; and NE = Not Evaluated. SEMARNAT status: A = Threatened; P = Endangered; Pr = Special Protection; and NS = No Status. See Alvarado-Diaz et al. (2013), Johnson et al. (2015a), and Mata-Silva et al. (2015) for explanations of the EVS, IUCN, and SEMARNAT rating systems. Taxon Sonora savagei** Sonora straminea** Sonora semiannulata Tantilla planiceps Trimorphodon lyrophanes Diadophis punctatus Hypsiglena catalinae** Hypsiglena chlorophaea Hypsiglena gularis** Hypsiglena marcosensis** Hypsiglena ochrorhynchus Hypsiglena slevini** Hydrophis platurus Rena boettgeri** Rena humilis Thamnophis elegans Thamnophis hammondii Thamnophis marcianus Thamnophis validus** Indotyphlops braminus*** Crotalus angelensis** Crotalus atrox Crotalus catalinensis** Crotalus cerastes Crotalus enyo** Crotalus helleri Crotalus lorenzoensis** Crotalus mitchellii** Crotalus polisi** Crotalus pyrrhus Crotalus ruber Crotalus thalassoporus** Caretta caretta Chelonia mydas Eretmochelys imbricata Lepidochelys olivacea Dermochelys coriacea Actinemys pallida Trachemys nebulosa** Trachemys scripta*** Kinosternon integrum*** Gopherus morafkai Apalone spinifera*** Amphib. Reptile Conserv. Distributional status P P P P FE E i E E E E E j E E NN E P P P P P NE3 : E E E E E E j E E P E E E3 3 3 3 3 E E 3 E 9 EB 3 3 3 7 Lo) lel 3 ie) 3 E BS EB FE E 3 E 9 9 9 9 9 3 E 3 NN NN NE NN Environmental Vulnerability Category (Score) ——_ Sl a IUCN categorization _—— | = ——<——_ = =. ss _——— LC NE IG LC LG. Le NE LC LC NE NE LC, LC NE LC LC L¢ LC Le, Le LC CR Le Le NE 1 LC NE NE iG NE LG. EN CR VU CR VU NE NE 88 SEMARNAT status Pr November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Table 9. Summary of the distributional status data for the herpetofaunal families in the Baja California Peninsula, Mexico. Number Distributional status Family of species Non-endemic (NE) Peninsular Endemic (CE) Non-native (NN) | SSS OL. ES eee ee a ee eee ee eae se er ipidac FE |) a ey ee See easy wep Rattidaes |e =e ee | ____* | Scaphiopodidae | 2 | Subtotal | | Plethodontidae [388 Le oa | ee = ea eee SSS SS SSS SSS SSS SSS SSS SSS TE i i ee EEE EE eee ) es Aniniellidaes eg —— biped ee ee | Crotaphytidae |S 8 | Eublepharidae | 3 | Gekkonidae | | dguanidae | | Phrynosomatidae | 30 | Phyllodactylidae |S | Scincidae | le Tiida | Xantusiidae | AT | Subtotal | 8B a a (a | | | | | Colubridae |S | Dipsadidae | jElapidags ee | Leptotyphlopidae | 2 a Natiicidae = | PA eee | Typhlopidae PS eiperidies ie eS | helontidae S| Se | Ease | Dermochelyidac | NP | Kinosternidae |S | Testudinidae |e | irionvatiidacs ef >=Es Lea ee a SE Lampropeltis catalinensis** Crotalus polisi** Masticophis barbouri** Crotalus thalassoporus** Rhinocheilus etheridgei** Sonora punctatissima** The 14 non-native species include five anurans (Smilisca Sonora savagei** baudinii, Xenopus laevis, Lithobates berlandieri, L. Hypsiglena catalinae** catesbeianus, and L. forreri), five lizards (Gehyra Hypsiglena gularis** mutilata, Hemidactylus frenatus, H. turcicus, Iguana Hypsiglena marcosensis** rhinolopha, and Sauromalus varius), one snake Crotalus angelensis** (Undotyphlops braminus), and three turtles (Trachemys Crotalus catalinensis** scripta, Kinosternon integrum, and Apalone spinifera). Crotalus lorenzoensis** The most widespread of these introduced species in Amphib. Reptile Conserv. 89 November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Mexico are Hemidactylus frenatus and Indotyphlops braminus (Gonzalez-Sanchez et al. 2021), and the most widely distributed of these species in the Baja California peninsula 1s Hemidactylus frenatus (Table 4). Principal Environmental Threats The main threat to the planet’s biodiversity is the combination of human population growth and _ the exploitation of natural resources. The human population requires resources to survive and grow, and _ these resources often are removed from the environment in unsustainable ways. This problem becomes larger as the human population grows. As threats to biodiversity are occurring worldwide, Mexico also is subject to many of these threats (Santos-Barrera et al. 2021), and the Baja California Peninsula is no exception. In this section we highlight the most significant problems we believe are affecting the conservation of amphibian and reptile populations in the Baya California Peninsula. Land conversion and habitat loss. In general, this threat is the major reason for biodiversity loss (Lecleére et al. 2020; Bellard et al. 2022), and specifically for amphibians and reptiles (Wake 1991; Gibbons et al. 2000; Bohm et al. 2013). In the Baja California Peninsula, a large part of this threat occurs mainly in coastal areas where the vegetation has been cleared, and the habitat has been lost as a result of tourism and housing development. For example, Aspidoscelis tigris and Dipsosaurus dorsalis were affected by habitat loss in the coastal sand dunes of San Felipe (Gatica-Colima 1998). This activity is more common in the Gulf of California due to the attractiveness of its beaches. Many of these mega-developments involve the clearing of large sections of natural land for the construction of buildings, golf courses, and marinas, which alter both the terrestrial and marine habitats, as well as coastal dunes and riparian areas (Rodriguez-Revelo et al. 2014b). In other regions, the expansion of agricultural areas also affects a large part of the habitats of many species throughout the peninsula (Fig. 25). Most of these crops are monocultures that serve as an ecological trap for organisms inhabiting nearby natural semiarid habitats (Rotem et al. 2013). One of the areas most affected by this activity is the Santo Domingo Valley in the Magdalena region of Baja California Sur. This area covers just over 260,000 ha and includes many isolated fragments of native vegetation that still contain native reptile species; nevertheless, extirpations have been detected, such as that of Urosaurus nigricaudus (Munguia-Vega et al. 2013). Perhaps this also is the case with other species of lizards and snakes in such agricultural areas as the Valley of Mexicali in northeastern Baja California, which covers about 280,000 ha. Another agricultural area with constant expansion is that between Colonet and San Quintin in northwestern Baja California, which has affected the coastal scrub vegetation and dune areas (Vanderplank % Fig. 25. Cleared land used for agriculture in San Quintin. This area was the habitat of Anniella geronimensis, and now is used by Driscoll’s to grow berries for export to the USA. The Riveroll Volcano is evident in the background, which is the habitat of Batrachoseps major and Ensatina schscholtzii. Agriculture and rock/sand mining activities continue to threaten these unique populations of salamanders. Photo by Jorge H. Valdez-Villavicencio. Amphib. Reptile Conserv. November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. et al. 2014a), thereby causing a reduction in the habitat of the endemic lizard Anniella geronimensis, as well as other endemic species (e.g., Aspidoscelis labialis and Sceloporus zosteromus). Mining concessions along the peninsula also imply the destruction of natural habitat, the excessive use of water, and habitat fragmentation due to the construction of access roads. In a short period between 2009 and 2010, 141 mining concessions were granted in the state of Baja California Sur, at least five of which are located in the Cape Region, which affected nearly 50,000 ha of habitat for many reptile species endemic to this region (Galina-Tessaro et al. 2015). Other mining activities, such as the extraction of sand from riparian areas that is used for construction and exported to the United States, have affected the habitat of the Arroyo Toad (Anaxyrus californicus) and the California Red-legged Frog (Rana draytonii) in northwestern Baja California; and both species are included in Mexico’s list of threatened species (Lovich et al. 2009). Water diversion and overuse. Aquatic habitats are threatened by the alteration of their physical or biotic structures based on the various ways humans use water supplies and the adjacent habitats (Figs. 26-28). The channelization of streams and excessive pumping of groundwater and surface water have the greatest effects on the aquatic habitats and their associated species (Jennings and Hayes 1994). Due to the aridity of the peninsula and the declining precipitation, evidently there is a diminution and a lack of recharge of the aquifers; therefore, the excessive use of water represents a strong threat to such freshwater species as amphibians and reptiles (e.g., Thamnophis and Actinemys). Notably, the extraction of water for domestic and agricultural consumption has affected the hydrology of the basins in northwestern Baja California. The excessive use of water for irrigation in agricultural areas is reducing the water levels in streams, and also causing the loss of ponds and areas with the historical presence of species such as Anaxyrus californicus, Rana draytonii, and Actinemys pallida (Peralta-Garcia et al. 2016; Valdez-Villavicencio et al., In Press). In most of the watersheds the surface water no longer occurs near the coast, and in some cases it is up to 20 km inland, due to the presence of a large number of pumps used to extract the groundwater and a network of surface pipes used to distribute water throughout the agricultural region between Colonet and San Quintin. Baja California Sur is no exception, as many of the aquifers there are overexploited, and with the scarcity of rainfall, the recharge of the aquifers is insufficient (Carrillo-Guerrero 2010; Troyo-Diéguez et al. 2010). Large amounts of water are destined for agricultural use. For example, 78% of the state’s water is destined for agricultural use and 15% for public establishments such as shopping areas and hotel complexes, whose numbers keep increasing with each passing year (Graciano 2013). This situation has caused a water deficit of 54% in the state’s aquifers (DOF 2020). Therefore, many of the streams and small oases no longer have the surface water which 1s vital for the survival of amphibians and reptiles (e.g., Trachemys nebulosa and Thamnophis hammondii). Invasive species. Introduced species are considered one of the main causes of amphibian and reptile Fig. 26. An aerial image of San Quintin agricultural valley, which surrounds Arroyo Santo Domingo. A large patch of wetland was present at the mouth of the stream, but became dry. Agricultural activities promote habitat loss through land clearing and water overextraction, which leads to soil salinization. Photo by Jorge H. Valdez-Villavicencio. Amphib. Reptile Conserv. 91 November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Fig. 27. Arroyo Santo Domingo, Baja California. A. Kilometers of water pipelines are used along Arroyo Santo Domingo to irrigate crops in the San Quintin valley; B. A well at Arroyo Santo Domingo. INEGI catalogued this arroyo as overexploited, but water extraction activities continue. Photo by Jorge H. Valdez-Villavicencio. Fig. 28. In the San Quintin Valley, fields are irrigated for agriculture. This area was the habitat of Anniella geronimensis and Aspidoscelis labialis, both endemic species in this area. Photo by Jorge H. Valdez-Villavicencio. declines (Blackburn et al. 2019; Cox et al. 2022), especially on island systems where invasive species are the main threat (Aguirre-Mufioz et al. 2016). Many of the alien species affect native species (Figs. 29-30), not only through direct predation, but also by competing for prey, modifying/eliminating habitat, and spreading disease (Holland 1994; Kats and Ferrer 2003; Bury et al 2012): Many species of mammals have been introduced on the islands associated with the Baja California Peninsula, some of which contribute to habitat loss and modification (e.g., goats), competition, and predation (e.g., rats, cats, and dogs). Cats prey directly on lizards and snakes, and could be their main threat on islands (Arnaud et al. 2008). Fortunately, many of these invasive species have been eradicated from several of these islands (Aguirre- Mufioz et al. 2016), but they still are present on some Amphib. Reptile Conserv. of the islands, which represents a threat to the unique species in these island habitats. In the northwestern region of the peninsula, a low abundance of amphibians, and the absence of Rana draytonii in particular, have been reported at sites where exotic species are present (Peralta-Garcia et al. 2016). The same situation has been observed with the pond turtle Actinemys pallida in the presence of Bullfrogs (Lithobates catesbeianus,; Valdez-Villavicencio et al., In Press). This same phenomenon occurs in many oases along the peninsula where various introduced species (e.g., Tilapia zilli, Procambarus clarkii, and L. catesbeianus) are found, which might be affecting the amphibian populations (Luja et al. 2016). In northeastern Baja California, the native anurans Incilius alvarius and Lithobates yavapaiensis perhaps have been extirpated due to the introductions of the Bullfrog and various fish species (Grismer 2002). November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. ¢ : eg 4 3 - 3 ‘: . at oe C yee — ia ott 4 = Fig. 29. Invasive Bullfrogs (Lithobates catesbeianus) at Valle de ~ & Guadalupe, Baja California. This species has dispersed throughout this valley due to growth in the wine industry, which usually includes reservoirs for irrigation or attractions for customers. Sunfish and Largemouth Bass have been released into these reservoirs, which threaten native amphibians and reptiles. Photo by Andrea Navarro-Tiznado. — he y. Fig. 30. An adult Hypsiglena ochrorhynchus was found in the stomach of a Bullfrog (Lithobates catesbeianus) at Rancho Madrigal, Ensenada, Baja California. Efforts to eradicate these frogs are being made at this site. Photo by Jorge H. Valdez-Villavicencio. In this same region, a reduction in the diversity of amphibian and reptile species also has been reported in disturbed sites of the Colorado River that have been invaded by exotic vegetation (Valdez-Villavicencio et al. 2021). Introduced plants can have serious negative effects on the environment and represent another threat to the survival of reptiles, since they can significantly reduce the richness and abundance of the insects that Amphib. Reptile Conserv. are the main food for many species of reptiles (Valentine et al. 2007; Schirmel et al. 2016). For example, the legless lizard Anniella stebbinsi appears to be affected by the invasion of ice plants (Carpobrotus edulis and Mesembryanthemum crystallinum) in the coastal dunes of northwestern Baja California (Manrriquez-Gomez et al. 2021). The same situation occurs in coastal dunes between San Quintin and El Socorro, where the November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Fig. 31. Livestock grazing at La Grulla, Sierra San Pedro Martir, Baja California, a high-elevation meadow at 2,250 m asl. The lack of livestock management affects local herpetofaunal populations, including those of Rana draytonii, Anaxyrus californicus, and Thamnophis elegans. Photo by Jorge H. Valdez-Villavicencio. Fig. 32. Livestock wandering along the side of Arroyo Valladares, Baja California. Photo by Anny P abundance of Anniella geronimensis has been observed to be much lower in patches with ice plants, when compared to patches of native vegetation (APG and JHVV, unpub. data). Invasive plants on island systems also might be affecting the survival of amphibians and reptiles. In recent years, the increase of non-native grass species has been observed on some Pacific islands. Introduced grasses have been shown to alter the structure and function of ecosystems, thereby causing negative impacts on native species (Garcia and Clusella-Trullas 2019), since they can reduce the availability of microhabitats, and also alter the thermal quality of the habitat (Carter et al. 2015; Schlesinger et al. 2020; Lara-Reséndiz et al. 2022). For example, a low abundance of Lampropeltis herrerae has been observed on Todos Santos Island, perhaps caused by introduced grasses (Pampa-Ramirez 2021), in addition to a low abundance of lizards. Amphib. Reptile Conserv. — ne eralta-Garcia. In other habitats such as oases, introduced plants represent a serious threat. The Rubber Vine (Cryptostegia grandiflora) is an aggressive invasive plant that has been introduced into at least 22 oases in Baja California Sur, and likely is affecting amphibians and reptiles associated with these unique habitats (Valentine et al. 2007: Rodriguez-Estrella et al. 2010). Livestock grazing. Livestock production is an important part of the economies of both states in the Baja California Peninsula, as the percentage of land use for cattle is over 50% (SEMARNAT 2003). The effects of ranching and livestock on wildlife need to be better documented (but see Figs. 31-32). However, overgrazing likely has led to a major problem in the meadows of northwestern Baja California, causing their disappearance and impacting riparian wildlife species, including amphibians such as Rana draytonii and Anaxyrus californicus (Mellink and November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. ees Fig. 33. A Southwestern Pond Turtle (Actinemys pallida) trampled Anny Peralta-Garcia. Contreras 2014; Peralta-Garcia et al. 2016). Remnant meadows might be affected by the water quality changes associated with grazing, leading to the demise of local amphibian populations (Smalling et al. 2021). In Baja California Sur, the tropical deciduous forest has been overgrazed for over 200 years; and during this time, grazing has changed the vegetational structure and composition of the Cape Region, thus affecting wildlife through habitat loss (Jaramillo 1994; Arriaga 2006). In general, habitats with a relatively complex structures support more diverse animal communities than those with simple structures, due to the provision of a greater range of available niches that can be exploited (Pianka 1966). Livestock grazing has been associated with a lower density and structure of the vegetation and seed production, soil compaction, the lowering of water filtration, increased erosion, as well as modification of the available oxygen, chemical composition, microorganisms, and fertility (Mellink and Contreras 2014; Jofré and Reading 2012). In the Baja California Peninsula, the abundance of three species of frogs and four species of lizards was lower in grazed areas than in ungrazed areas (Arguelles-Méndez et al. 1996; Romero- Schmidt et al. 1994; Romero-Schmidt and Ortega-Rubio 1999). The differences were attributed to a reduction in the protective cover from predators and food availability for insects. Furthermore, cattle might have a direct effect by disturbing individuals or even crushing them as they wander in their habitat. Cattle grazing had a detrimental effect on lizard populations. For example, Busack and Bury (1974) found negative effects on Gambelia Amphib. Reptile Conserv. 95 oY * f by cattle along Arroyo San Vicente, Baja California. Photo by wislizenii and Phrynosoma platyrhinos (two species found in Baja California), because grazing had altered their preferred habitats and the availability of food. Dead individuals of Actinemys pallida have been found on stream banks (Fig. 33), showing signs of being crushed by livestock (APG and JHVV, pers. obs.). Grazing and trampling led to an overall decline in reptile population abundance, changes in reptile species composition, and reduced reptile diversity in the majority of the habitat types where it has been studied (Jofré and Reading 2012). However, more research is needed to understand the effect of grazing on amphibians and reptiles in the Baja California Peninsula. Illegal trade. Currently, many species of amphibians and reptiles that are rare or endemic, or that inhabit particular areas such as islands, are attractive to collectors and often are illegally collected and traded, which can lead to over- collecting, consequently affecting their survival (Auliya et al. 2016; Marshall et al. 2020). The herpetofauna of the Baja California Peninsula is known to be subject to the illegal collecting and trafficking of species (Mellink 1995; Grismer 2002). Many insular endemic species are subject to illegal collection and trade. The Baja California associated islands harbor numerous endemic species that are subject to the pet trade, especially rattlesnakes (Pliego-Sanchez et al. 2021) and the Todos Santos Island Kingsnake (Lampropeltis herrerae). Collectors turning over rocks and snake traps have been reported on Isla Todos Santos (Mellink 1995). In 2007, APG and JHVV also found several abandoned traps, and recently Pampa- November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Fig. 34. Car and motorcycle tracks seen in the vicinity of turtle nesting sites at Punta Arena, north of Cabo Pulmo, Baja California Sur. These activities remain a common practice along these areas. Photo by Alan Harper. Ramirez (2021) reported finding three more abandoned traps. However, we do not know whether this species presently continues to be attractive for illegal collection. The search for and illegal collection of reptiles also has led to the degradation of the habitat of rock-dwelling species (Goode et al. 2004, 2005), such as the rocky habitat of the snakes Lampropeltis multifasciata and Lichanura trivirgata, and rock-dwelling lizards (e.g., Petrosaurus and Sauromalus). Reports of the illegal removal of large numbers of the endemic Petrosaurus thalassinus in the Cape Region of Baja California Sur have been published (Grismer 2002; Lovich et al. 2009). Recently, we have seen social media posts of people purchasing individuals of Bipes biporus and Phrynosoma species from Baja California. This practice encourages the local people to illegally (sometimes unknowingly) collect reptile species to sell to these types of buyers. Off-road activities. Off-road vehicle driving (Figs. 34-35) is a common outdoor recreational activity in the Baja California Peninsula (Gaeta-Verdin 2020). The use of these types of vehicles (quads, buggies, racers, etc.) has increased significantly in recent years, but there are few regulations on how and where these recreational activities can be undertaken. Today, these activities are a common tourist attraction along the coasts of the peninsula, where rental services for these vehicles are offered. Few studies in Baja California have evaluated whether these activities affect biodiversity, but studies elsewhere have shown the negative effects on reptiles in desert habitats due to the loss of vegetational cover, a reduction in the abundance of invertebrates, and the disturbance of the daily activities of reptiles (Busack and Bury 1974; Bury et al. 1977). One example is the Amphib. Reptile Conserv. alteration of the coastal dunes in northwestern Baja California (El Descanso dunes, and La Lagunita in Ensenada), where the vegetation has been lost completely due to the use of all-terrain vehicles (Rodriguez-Revelo et al. 2014a), and it surely has affected species such as Anniella stebbinsii and Sceloporus zosteromus. These types of activities also could be affecting other species, including Phrynosoma mcallii, Uma notata, and Crotalus cerastes (included in NOM-059), as they may be run over by off-road vehicles passing through Laguna Salada or the Cuervitos-Algodones dunes in Mexicali. The use of these vehicles in riparian areas also is a common activity, sometimes directly in the streambeds, where they affect freshwater turtles and tadpoles or amphibian egg masses. Baja California Sur is no exception. Due to the increasing amount of tourism, numerous places now offer these types of recreational activities, and mainly in coastal areas. Sea turtle nesting areas also have been affected by tourists or local people who do not respect the posted signs, and use motorcycles and cars to enter the sand dunes where turtles nest (Vanderplank et al. 2014b). High-speed off-road vehicle races are held in Baja California each year (e.g., Baja 1,000, 500, 400, and 250 mile races). Nearly 300 vehicles participate in the Baja 1,000 and travel along the dirt roads and trails along the peninsula, and even through natural protected areas. These vehicles likely are causing irreparable damage to the arid and semi-arid environments of the Baja California peninsula, since constant traffic compacts the soil, destroys the vegetation, and promotes erosion (Gaeta-Verdin 2020). Given the characteristics of these vehicles, Scaphiopus couchii and Spea hammondii burrows can be susceptible to the disturbance caused by the vehicles that generate noise and produce vibrations November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Fig. 35. Well-traveled road along a streambed at Cafion El Alamo, Arroyo Las Palmas, Baja California. Several species of amphibians and reptiles, including Southwestern Pond Turtles (Actinemys pallida), inhabit this stream. Photo by Anny Peralta-Garcia. similar to rain, inducing their emergence under highly unfavorable conditions (hot and dry) that would be fatal for adults (Jennings and Hayes 1994). Another problem with this activity is that the routes are not permanent, and every year they change and affect different areas. The traffic caused by local vehicles also is a problem, as thousands of spectators also travel along the dirt roads to watch the races at different points, and they surely run over a large number of lizards and snakes, in addition to the problem of improperly disposing of their garbage (Gaeta- Verdin 2020). Infectious diseases. Infectious diseases are considered one of the main threats to amphibians worldwide, resulting in the declines and local extinctions of many species. One of the main diseases is chytridiomycosis, an infectious disease caused by the pathogenic fungi Batrachochytrium Amphib. Reptile Conserv. dendrobatidis and B. salamandrivorans (Skerratt et al. 2007; Adams et al. 2022). For reptiles, some infectious diseases also have emerged and affected wild populations through disease transmission, such as the one in snakes caused by the fungus Ophidiomyces ophiodiicola, and the turtle shell disease caused by Emydomyces testavorans (Haynes et al. 2021; Lambert et al. 2021). Only a few studies in the Baja California Peninsula have investigated diseases in amphibians, and they have focused on B. dendrobatidis (Bd). Bd has been recorded in both states, with museum records detecting the pathogen as early as 1932 in Baja California (Adams et al. 2022). In the northern state, this fungus has been detected in Anaxyrus californicus, A. boreas, Pseudacris cadaverina, P. hypochondriaca, Rana draytonii, and the exotic Lithobates catesbeianus, with only Xenopus laevis testing negative (Peralta-Garcia et al. 2018). In November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Baja California Sur, Bd also has been detected in L. catesbeianus and P. hypochondriaca (Luja et al. 2012). Bd-positive sites occurred at elevations ranging from sea level to 2,070 m asl. The presence of the non-native American Bullfrog (Lithobates catesbeianus), a competent Bd vector and reservoir host (Schloegel et al. 2012; Adams et al. 2017), has been associated with higher Bd prevalence in native anurans in Baja California (Luja et al. 2012). Increased Bd prevalence in Baja California also has been observed at higher elevations, and with greater remoteness from urban areas and agricultural land (Peralta-Garcia et al. 2018). Adams et al. (2022) found an overall Bd prevalence of 68%, with species being an important predictor of pathogen prevalence and burden (load) both across and within sites. Species distribution models of Bd predict high suitability for the pathogen in northwestern Baja California, based on environmental factors (Bolom- Huet et al. 2019). Infections higher than >10,000 ZE (where ZE is a measure of infection intensity) have been observed in Baja California frogs, particularly in A. boreas and R. draytonii, with a higher prevalence at higher elevation sites (Peralta-Garcia et al. 2018; Adams et al. 2022), prompting the need for further investigations of Bd in this region. No diseases have been detected for reptiles, but follow-up studies are important because some diseases have been detected in snakes and turtles in the neighboring state of California (Haynes et al. 2021; Lambert et al. 2021). Climate change. Climate change has become one of the main threats to biodiversity (Pereira et al. 2010; Bellard et al. 2012). Some of the main effects of climate change are changes in temperatures that can accelerate the loss and degradation of habitat, promote changes in the abundance and structure of communities, and alter the distribution of species, in addition to accelerating the extinction of species at different scales (Bellard et al. 2012). In the case of reptiles, many studies have examined the impact of climate change and indicate severe effects on these organisms (Sinervo et al. 2010; Meiri et al. 2013). Climate change represents a threat for most reptiles since it can reduce their hours of activity, thereby causing the alterations in many of their physiological processes, reproduction, and feeding (Sinervo et al. 2010). In the Baja California Peninsula, species with restricted distributions or species with fossorial habits are the ones that primarily will be affected. For example, climate change effect models show that Anniella geronimensis and Bipes biporus would experience negative impacts on their distributions (Lara-Reséndiz et al. 2020). Conversely, some thermophilic and widely distributed species (e.g., Dipsosaurus dorsalis) might not be as threatened by climate change (Lara-Reséndiz et al. 2019). Fossorial species (such as Anniella geronimensis and Bipes biporus) may also be impacted under projected Amphib. Reptile Conserv. climate change scenarios, in which reduced dispersal and mobility may be coupled with reduced suitable habitat (Lara-Reséndiz et al. 2020). The effects of climate change could present a severe threat to insular systems, since amphibians and reptiles obviously cannot expand or modify their distributions to compensate for its effects. Among terrestrial vertebrates on islands, reports show that amphibians and reptiles could be the most affected by climate change. Estimates indicate that many species could lose close to 50% of their distribution ranges (Ureta et al. 2018). In addition to this factor, the rise in sea level due to the effects of climate change would affect many of the reptiles on islands (Bellard et al. 2013; Pliego-Sanchez et al. 2021). Although the effects of climate change on amphibians in the Baja California Peninsula have not been evaluated, we realize that changes in temperature on a global scale also will have strong effects on amphibians. These changes could affect reproduction and hibernation periods, as well as their ability to find food, alter pathogen-host dynamics, lead to increased stress from UV radiation, as well as alterations in reproduction and hibernation periods (Blaustein et al. 2010). Species that inhabit elevations above 500 m are expected to lose a significant part of their climatically suitable area (Alves-Ferreira et al. 2022). Conversely, species that inhabit arid environments tend to expand their ranges in response to climate change. This result can be explained by the environmental characteristics of these habitats, which tend to have extreme seasonal climates with well-defined periods of drought and rain (Alves-Ferreira et al. 2022). Although the Cape Region in Baja California Sur is not particularly diverse in amphibian species, some predictions also indicate a reduction in the number of species that inhabit tropical dry forest (Ballesteros-Barrera et al. 2022). Conservation Status We used the three systems of conservation assessment that were used in the previous entries in the Mexican Conservation Series (MCS; see above). These systems are SEMARNAT (2019), the IUCN Red List (http:// iucnredlist.org), and the EVS (Wilson et al. 2013a,b). Based on the features of the Baja California Peninsula system, we updated the assessments from these three systems as necessary. The SEMARNAT List of Threatened Species The Mexican Federal Government designates threatened species on the NOM-059-SEMARNAT-2010 listing. Species are classified through a risk assessment method developed by the Secretaria del Medio Ambiente y Recursos Naturales, with the last update published in 2019 (SEMARNAT 2019). The available ratings from this list are provided in Table 7 and summarized in November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Table 10. SEMARNAT categorizations for the herpetofaunal species in the Baja California Peninsula, Mexico, arranged by family. Non-native species are excluded. Number of species 142 158 I i eS) — = Table 10, and non-designated species are not included on the NOM-059-SEMARNAT-2010 list. This system utilizes three categories of assessment: endangered (P), threatened (A), and under special protection (Pr), and non-designated species are indicated here by using a “no status” (NS) category. The data in Table 10 show that 85 (53.8%) of the 158 native species inhabiting the Baja California Peninsula and its adjacent islands are included in NOM-059 SEMARNAT, whereas 73 (46.2%) are not included, which could indicate either that they have not been assessed or are not considered as threatened. In all MCS studies, the question always arises as to whether any bias is shown toward the conservation assessments of endemic species as opposed to non- endemic species using the SEMARNAT system, inasmuch as the majority of the species in a given area examined, in this case the Baja California Peninsula and its adjacent islands, remain unassessed. In order to answer this question, the pertinent data are shown in Table 11. These data show that about one-half of the Amphib. Reptile Conserv. SEMARNAT categorization Special Endangered (P) | Threatened (A) Protection (Pr) No Status “ non-endemic species (42, or 51.9% of 81 total species) remain unassessed, while a slightly lower number were unassessed for the peninsular endemics (31, or 40.3% of 77 total species). For the total native herpetofauna, 74, or 46.8% of 158 species, remain unassessed. Thus, no clear bias is apparent toward the peninsular endemics. The IUCN System The implementation of the system of conservation assessment established by the International Union for the Conservation of Nature has not kept pace with new species descriptions and ongoing taxonomic research. In fact, this 1s why the EVS system was developed—e., to assist in the development of conservation strategies when the IUCN system has not been applied. The data for the IUCN categorizations are shown in Table 8 and summarized in Table 12. Of the 158 native species in the herpetofauna of the Baja California Peninsula, 130 (82.3%) have been evaluated under the IUCN system (Table 12). Of these 99 November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 11. Comparison of SEMARNAT and distributional categorizations for the Baja California Peninsula herpetofauna. Non- native species are excluded. SEMARNAT category Endangered (P) Threatened (A) we Eee 5 ae Distributional category 130 species, 15 (11.5%) have been placed in one of the three “threat categories,” including three in the CR category, four in the EN category, and eight in the VU category. The three species in the CR category are the snake Crotalus catalinensis, a peninsular endemic species; and the turtles Eretmochelys imbricata and Dermochelys_ coriacea, both non-endemic marine species. The four species in the EN category are the anuran Anaxyrus californicus, a non-endemic species; the lizards Anniella geronimensis and Sauromalus hispidus, both peninsular endemic species; and the turtle Chelonia mydas, a non-endemic marine species. The eight VU species are the lizards Sauromalus klauberi, Uta encantadae, U. lowei, U. tumidarostra, Aspidoscelis catalinensis, and A. labialis, all peninsular endemic species, and the turtles Lepidochelys olivacea and Actinemys pallida, both non-endemic species. Of the 115 species placed in the “lower risk categories” Table 12. IUCN Red List categorizations for herpetofaunal families in the Baja California Peninsula, Mexico. Non-native species are excluded. Shaded columns to the left are the “threat categories,” and those to the right are the categories for which too little information on conservation status exists to allow the taxa to be placed in any other IUCN category, or they have not been evaluated. Critically Endangered UTES Family , species | 13 1 4 8 3 2 4 1 2 158 158 6 30 3) 3 7 1 29 7 1 2 5 14 Amphib. Reptile Conserv. Near Vulnerable | +) +-eatened IUCN Red List categorization Least Concern 5 Data Not Deficient Evaluated = N N (EES ¢ November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. (NT and LC), only seven are in the NT category, with the remaining 108 species in the LC category (Table 12). The seven NT species are the anurans Rana draytonii, Rana boylii, and Spea hammondii, all non-endemic species, and the lizards are Sauromalus slevini, Phrynosoma mcallii, Uma notata, and Phyllodactylus unctus, which include two peninsular endemic species and two non-endemic species. The 108 LC species make up 68.4% of the 158 native species in the Baja California Peninsula. Finally, 28 species have not been assessed using the [UCN system, including three allocated to the DD category. These 28 Species comprise 17.7% of the native species, and their assessments using the EVS system are given below. The EVS System As discussed in all the earlier MCS _ studies, the Environmental Vulnerability System (EVS) initially was developed for use in the conservation evaluation of the herpetofauna in the country of Honduras (Wilson and McCranie 2004). Those authors created this system as a means for determining the conservation status of the members of a herpetofauna whose knowledge was insufficient when using the IUCN system. When the work of the MCS began in 2013, it was evident that the EVS could be applied to the Mexican herpetofauna just as easily as it had for the Honduran herpetofauna. Subsequently, this system has been employed in all of the MCS studies to date (see above), including the present one. Thus, the EVS values for the 152 native non-marine species of the Baja California Peninsula are shown in Table 8 and summarized in Table 13. The EVS values range from 3 to 20. The most frequent values (applied to 10 or more species) are 9 (11), 10 (10), 11 (15), 12 (14), 14 (15), 15 (15), 16 (18), and 17 (29). Collectively, these eight values were applied to 127 of the 152 native non-marine species (83.6%). The lowest score of 3 was ascertained for two anuran species (Scaphiopus couchii and Anaxyrus punctatus) and the highest score of 20 was assigned to the Todos Santos Island Kingsnake (Lampropeltis herrerae). The EVS scores are grouped into three categories Table 13. Environmental Vulnerability Scores (EVS) for the herpetofaunal species in the Baja California Peninsula, Mexico, arranged by family. The shaded area to the left encompasses low vulnerability scores, and the one to the right high vulnerability scores. Non-native species are excluded. Number Environmental Vulnerability Scores | Environmental Vulnerability Scores aS ee eee | | | | Bufonidae | 6 | te | 1 | ~ | ~] 21 [=| Se oo | Ranidae | 3 eee) — |) | 2 | — (ee | Scaphiopodidac | (AGMA eee) — | 1 | — | — [ESS ee ee Subtotal | 3) Ens See i |: |? | ahaa | Plethodontidac || 3 (a eee ee) — | — | — | — [Se | Subtotal | 3S ee) — | — | — | — [ee | Total | eee — | — | — | — eee |Anguidee | 5S se eee! | |—|—|- Rn |Amiclide || 2. See —| | |—|- Rn Eee | Bipedidae | ees —~ | — | — | — [ee L-Crotaphycidae [S| Se emcees — {1 | 1 | — Re —_ oo == Ee ee || ||: |-|> Be Family | Phyllodactylidac | S(t ee ee)! | — | — | — [ee | Scincidae Scincidae | 3 =a Ea) | ||!) aaa aa ee | Teiidac |S eee eee || — | — | — [ee | Xantusiidac | 4 ee eee) — | 2 | — | — [SR eee | Subtotal =| 8 TOL OT at ToT etait ato {oto {4 {se {o [sii] olio | | Charinidae | eee | | — | — | — eee | Colubridac | 2 ee —|2 |: |) aaa | Dipsadidac | 7 eee 2 | | — | — ee | Leptotyphlopidac | (INN re eet OMe) — | — | — | — [RSS eee | Natricidac | 4 eee) — | | | — ee | Viperidac | 12 ees — | — | 1 | 2 ee | Subtotal | ST OT tT tT at Tat Ts To] st 4 tof] sts | 4 ts [3 [2 {4 [| | Emydidae | Ee ee) — | — | — | ee | Testudinidac |e ee ee ee) — | — | — | — [ee | Subtotal | TOT Of of of of ofofololfoltifol2z{o{ofol{o{o | Total | GS | OT | 2 Ta] 3 | © [0] 9 | to] 2] 8 | 13 | 15 | 18 | 20] 2 | 5 | 0 | Sams total _|__152__ [RAISES RSAC 10 | 1s | 14 | 9 SIRS ES | Categorytotal | 52H Amphib. Reptile Conserv. 101 November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 14. Comparison of Environmental Vulnerability Scores (EVS) and IUCN categorizations for members of the herpetofauna of the Baja California Peninsula, Mexico. Non-native species are excluded. The shaded area at the top encompasses low vulnerability scores, and the one at the bottom high vulnerability scores. IUCN category Critically iRndanoered Vulnerable Near Least Data Not Total a 8 a ss a = of low, medium, and high vulnerability. As a result, the summary values (Table 13) increase from low vulnerability (28 species) to medium vulnerability (48 species), and then to high vulnerability (76 species). Typically, this pattern is characteristic of herpetofaunas containing more endemic than non-endemic species, although this is not the case with the herpetofauna of the Baja California Peninsula, in which there are 83 non- endemic species and 75 endemic species. However, this pattern could be due to the high level of island endemics that have high vulnerability values. The numbers of species in the herpetofauna of the Baja California Peninsula for each IUCN/EVS score combination are shown in Table 14. These data illustrate that although both systems agree on the low vulnerability category and Least Concern category for 26 species (17%), they differ in the remaining categories, as only 11 of the 76 high vulnerability species (14.5%) are placed into one of the three IUCN “threat categories” (CR, EN, or VU). As was found in all other MCS studies, the results of the application of the IUCN and EVS systems of conservation assessment do not correspond well with one another. Of the 152 species that can be assessed by both the IUCN and EVS systems, only three have been allocated to the IUCN DD category (Table 15). These are three peninsular endemic snake species (Lampropeltis catalinensis, Masticophis barbouri, and Rhinocheilus etheridgei), which have respective EVS scores of 18, 17, and 17. Based on the arguments presented in previous MCS studies, we suggest that once these species are evaluated by the IUCN, they should be relegated to the CR (Lampropeltis catalinensis) and EN (Rhinocheilus etheridgei and Masticophis barbouri) categories. Twenty-five species (15.8% of the 158 native species) have not been evaluated by the IUCN system. These 25 species include seven lizards, 16 snakes, and two turtles. The majority of these species (15 of 25, or 60.0%) are peninsular endemics, and the remainder (10, or 40%) are non-endemics. These species are allocated to the three EVS categories of vulnerability as follows: two low (8.0%); seven medium (28.0%); and 16 high (64.0%). Based on the vulnerability values, species with an EVS of 17 and greater might be placed in the CR category; Table 15. Environmental Vulnerability Scores (EVS) for members of the herpetofauna of the Baja California Peninsula, Mexico, that are allocated to the IUCN Data Deficient category. ** = peninsular endemic. Environmental Vulnerability Score (EVS) Taxon Geographic Ecological Reproductive mode/ Total —=—- ——V——re —e—eeeo of — score | Lampropeltis catalinensis** | catalinensis | Lampropeltis catalinensis** | ig Masticophis barbouri** Se a rs Rhinocheilusetheridgei** | 6 EBT Amphib. Reptile Conserv. November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. No. 1. Anaxyrus californicus (Camp, 1915). The Arroyo Toad is found “west of the deserts in southern California (USA) near Santa Margarita in San Luis Obispo County south into northern Baja California (Mexico), at least as far south as Arroyo San Simon, just south of San Quintin” (Frost 2022). This individual came from Rancho Meling, Sierra San Pedro Martir, in the municipality of San Quintin. In this study its EVS was estimated as 11, placing it in the upper portion of the medium vulnerability category. The IUCN has assessed its conservation status as Endangered (EN) and SEMARNAT lists it as Threatened (A). Photo by Ivan Parr. No. 3. Rana draytonii (Baird and Girard, 1852). The California Red-legged Frog is distributed from “Mendocino County (California, USA) south along the Pacific coast of the USA to the vicinity of Arroyo Santo Domingo in northern Baja California (Mexico)” (Frost 2022). This individual was found at Rancho Meling, Sierra San Pedro Martir, in the municipality of San Quintin. In this study its EVS was indicated as 11, placing it in the medium vulnerability category. The IUCN evaluated its status as Near Threatened (NT), and this species is listed by SEMARNAT as in danger of extinction (P). Photo by J.A. Soriano. Amphib. Reptile Conserv. 103 No. 2. Pseudacris hypochondriaca (Hallowell, 1854). The Baja California Chorus Frog ranges from “southern California, western and southwestern Nevada, and adjacent northwestern Arizona (Mohave County) south to the southern tip of Baja California peninsula, Mexico” (Frost 2022). This individual was found at Rancho Meling, Sierra San Pedro Martir, in the municipality of San Quintin. In this study its EVS was determined as 9, placing it at the upper limit of the low vulnerability category. This species has not been evaluated by the IUCN or SEMARNAT. Photo by Jorge H. Valdez- Villavicencio. Salamander ranges “from sea level to near 1,270 m in elevation and ranges from Eureka in Humboldt County, California, south through the Coast Ranges, terminating near Valle Santo Tomas in northwestern Baja California. It also occurs on the Coronado Norte island” (Grismer 2002: 56). This individual came from La Mision, Baja California, in the municipality of Ensenada. In this study its EVS is listed as 14, placing it at the lower limit of the high vulnerability category. The IUCN judged its conservation status as Least Concern (LC), and SEMARNAT lists it as a species of Special Protection (Pr). Photo by Jorge H. Valdez-Villavicencio. November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 16. Environmental Vulnerability Scores (EVS) for members of the herpetofauna of the Baja California Peninsula, Mexico, that are currently not evaluated (NE) by the IUCN. Non-native taxa are excluded. ** = peninsular endemic. Environmental Vulnerability Score (EVS) Taxon Geographic Ecological Reproductive mode/ Total distribution distribution Degree of persecution score 1 11 [Amnietiasebbinsis =i [Hxpsiglona ochroriynctus [4 [Rena boenger™ «dS [Crotalus heltes — | 15 Phyllodactylus xanti** i Plestiodon gilberti Nn Wo — oS) Plestiodon lagunensis** Plestiodon skiltonianus Aspidoscelis canus** Aspidoscelis carmenensis** Aspidoscelis celeripes** epee — STN] oO — Aspidoscelis ceralbensis** Aspidoscelis danheimae** Oo Aspidoscelis espiritensis** —~ Aspidoscelis franciscensis** j-) Aspidoscelis hyperythrus — Oo — Aspidoscelis pictus** Aspidoscelis tigris — — — — —s — — — Nn — Xantusia henshawi -) Lichanura trivirgata Arizona elegans — Arizona pacata** Bogertophis rosaliae Lampropeltis californiae N >) Lampropeltis herrerae** Masticophis aurigulus** Masticophis flagellum Masticophis lateralis — Phyllorhynchus decurtatus Pituophis catenifer ~“ Pituophis insulanus** N eS) Pituophis vertebralis** Rhinocheilus lecontei Salvadora hexalepis Sonora savagei** Sonora semiannulata Tantilla planiceps Trimorphodon lyrophanes Diadophis punctatus Hypsiglena chlorophaea Oo Hypsiglena gularis** is Hypsiglena slevini** Rena humilis | lela — — —[—}_) im >) — Oo — Thamnophis elegans Thamnophis hammondii Thamnophis marcianus Thamnophis validus** Crotalus angelensis** CoOoTr Crotalus atrox Crotalus cerastes Uo Crotalus enyo** Crotalus lorenzoensis** Crotalus mitchellii** Crotalus ruber — en oo WTO | Phyllodactylusxani** | ST | Plestiodongilberti | | Plestiodonlagunensis** |S | Plestiodon skiltonianus | 3S | Aspidosceliscanus** | | Aspidoscelis carmenensis** | 6B | Aspidoscelis celeripes** | SE | Aspidoscelis ceralbensis** | OB | Aspidoscelisdanheimae** |_| 8 | Aspidoscelis espiritensis** |S | Aspidoscelis franciscensis** | | Aspidoscelishyperythrus | 2S | Aspidoscelispictus** | OE | Aspidoscelistigris | | Xantusiahenshawi | Lichanuratrivirgata | | Arizonaelegans | | Arizonapacata**® TS | Bogertophisrosaliae | | Lampropeltis californiae | A | Lampropeltisherrerae** | OE | Masticophisaurigulus** |S | Masticophis flagellum | 8 | Masticophislaterais | 3S | Phyllorhynchusdecurtaus | AES | Pituophiscatenifer | Pituophis insulanus®* | Pituophisvertebralis**® |S | Rhinocheiluslecomei_ | tT 8 | Salvadorahexalepis | | Sonorasavager** | Sonorasemiannulata | | Tantillaplaniceps | 8 | Trimorphodon lyrophanes | | Diadophis punctatus | Hypsigiena chlorophaca | 3S | Hypsiglenagularis®* | 8 | Hypsigienaslevini®* TS | Renahumilis 8 | Thamnophiselegans | | Thamnophishammondii | AES | Thamnophis marcianus |S | Thamnophisvalidus** |S | Crotalusangelensis** | OT | Crotalusatrox, 8 | Crotalus cerastes TT | Crotalusenyo*® TS 8 | Crotaluslorenzoensis®* |B | Crotalusruber Amphib. Reptile Conserv. 107 November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula at No. 9. Anniella geronimensis Shaw, 1940. The Baja California Legless Lizard “ranges along the coastal aeolian dune regions of northwestern Baja California, from approximately 6 km north of Colonia Guerrero south to just south of Punta Baja at the northern edge of Bahia El Rosario. It is also known from the Pacific islands of San Geronimo and San Martin.” (Grismer 2002: 242). This individual was encountered at San Quintin, Baja California, in the municipality of San Quintin. In this study its EVS is noted as 14, placing it at the lower limit of the high vulnerability category. The IUCN judged its conservation status as Endangered (EN), and SEMARNAT lists it as a species of Special Protection (Pr). Photo by Jorge H. Valdez-Villavicencio. No. 11. Crotaphytus insularis Van Denburgh and Slevin, 1921. The Isla Angel de la Guarda Collared Lizard is endemic to Isla Angel de la Guarda in the Gulf of California in the municipality of Mexicali (Grismer 2002). Its EVS was assessed as 16, placing it in the middle of the high vulnerability category. The IUCN evaluated its conservation status as Least Concern (LC), but this lizard is not listed by SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. Amphib. Reptile Conserv. ts & le y q) 2 | es pt ey ~~ 2 he a — ~G ~ Bp No. 10. Bipes biporus (Cope, 1894). The Five-toed Worm Lizard is distributed “throughout the western portion of the southern half of Baja California peninsula, west of the Peninsular Ranges, from approximately 17 km north of Jesus Maria, where the Sierra Columbia contacts the Pacific coast, south to Todos Santos (Mahrdt et al. 2022)...At the Isthmus of La Paz, its distribution extends east across the low, sandy flats and contacts the Gulf coast at Bahia de La Paz” (Grismer 2002: 254). It also occurs on the Pacific island of Magdalena (Peralta-Garcia et al. 2007). This individual was found at La Paz, Baja California Sur, in the municipality of La Paz. In this study its EVS was determined as 19, placing it near the higher limit of the high vulnerability category. The IUCN assessed its conservation status as Least Concern (LC), and SEMARNAT designates it as a species of Special Protection (Pr). Photo by Jorge H. Valdez-Villavicencio. No. 12. Gambelia copeii (Yarrow, 1882). The Baja California Leopard Lizard “is endemic to Baja California peninsula and ranges from extreme southern San Diego County, California, south to at least Todos Santos on the west coast of the Cape Region” (Grismer 2002: 114). This individual came from 20 km NW of San Juanico, Baja California Sur, in the municipality of Comondu. In this study its EVS was estimated as 11, placing it in the lower portion of the medium vulnerability category. The IUCN judged its conservation status as Least Concern (LC), but this species is not listed by SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. No. 13. Coleonyx switaki (Murphy, 1974). Switak’s Banded Gecko “ranges along the desert foothills of the Peninsular Ranges from at least northern San Diego County, California, south to just north of Santa Rosalia” (Grismer 2002: 199) in Baja California Sur. This individual was located in San Ignacio, Baja California Sur, in the municipality of Mulegé. In this study its EVS is noted as 10, placing it at the lower limit of the medium vulnerability category. The IUCN determined its conservation status as Least Concern (LC), but this gecko is not listed by SEMARNAT. Photo by Tim Warfel. No. 14. Ctenosaura hemilopha (Cope, 1863). The Cape Spiny- tailed Iguana “ranges from near Loreto south along the Sierra la Giganta to the west coast near Arroyo Seco and through the Cape Region...In the Gulf of California, C. hemilopha is known only from Isla Cerralvo...” (Grismer 2002: 117). This individual came from Sierra La Gata, Baja California Sur, in the municipality of La Paz. In this study its EVS was calculated as 16, placing it in the middle portion of the high vulnerability category. The IUCN determined its conservation status as Least Concern (LC), but SEMARNAT considers it to be a species of Special Protection (Pr). Photo by Jorge H. Valdez-Villavicencio. No. 15. Sauromalus ae tanta 1891. The Spiny Chuckwalla “is known from the Gulf islands of Angel de la Guarda, Cabeza de Caballo, Flecha, Granito, La Ventana, Mejia, Piojo, Pond, San Lorenzo Norte, San Lorenzo Sur, and Smith” (Grismer 2002: 128). This individual was found on Isla Angel de la Guarda, Baja California, in the municipality of Mexicali. In this study its EVS was determined as 14, placing it at the lower limit of the high vulnerability category. The IUCN assessed its conservation status as Endangered (EN), and SEMARNAT noted it as Threatened (A). Photo by Jorge H. Valdez-Villavicencio. Amphib. Reptile Conserv. PRES Fe ta eke No. 16. Callisaurus draconoides Blainville, 1835. The Zebra- tailed Lizard “ranges throughout the Sonoran and Mojave deserts of the southwestern United States and northern Mexico south to southern Sinaloa and all the arid regions of Baja California” (Grismer 2002: 136). This individual was encountered at Guerrero Negro, Baja California Sur, in the municipality of Mulegé. In this study its EVS is listed as 12, placing it in the upper portion of the medium vulnerability category. The IUCN assessed its conservation status as Least Concern (LC), and SEMARNAT designates it as Threatened (A). Photo by Alan Harper. November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 18. Number of herpetofaunal species in three distributional status categories among the 10 physiographic regions of the Baja California Peninsula, Mexico. Rank was determined by adding the state and country endemics. salamanders Aneides lugubris and Batrachoseps major, both with an EVS of 14 (Table 8). In Baja California, the major threats to amphibians are wetland habitat loss and the presence of exotic species. If regional specific criteria were considered in future EVS_ evaluations, vulnerability values for amphibians in this region likely would be higher and correspond more closely with those from other parts of Mexico. Relative Herpetofaunal Priority Johnson et al. (2015) introduced the concept of Relative Herpetofaunal Priority (RHP) as a simple means for measuring the relative importance of the herpetofaunal species in any geographic segment (e.g., state or physiographic region). Ascertaining the RHP involves the employment of two methods: (1) calculating the absolute number of state, country, or regional endemic species as they relate to the entire regional herpetofauna, and (2) calculating the absolute number of high EVS category species in the entire regional herpetofauna. The pertinent data for these two methods are shown in Tables 18 and 19. Based on the relative number of peninsular endemics (Table 18), the 1* rank is held by the GIR with 50 peninsular endemics from a total of 84 species (59.5%). The remaining ranks are as follows: 2" is ATR (26 of 65 species, 40.0%); 3 is CGCR (24 of 62 species, 38.7%); 4" is SLLR (21 of 41 species, 51.2%); 5" (held by two regions) is MR (18 of 51 species, 35.3%) and VR (18 of 64 species, 28.1%); 6" is PIR (15 of 47 species, 31.9%); 7" is CR (eight of 60 species, 13.3%); 8 is LC VR (five of 68 species, 7.4%); and 9" is BCCFR (0 of 27 species, 0.0%). Clearly, the highest herpetofaunal priority is associated with the Gulf Islands Region. This region consists of all the islands lying off the Baja California Peninsula and within the Gulf of California, excluding those more closely associated with the state of Sonora on the eastern side of the Gulf. Collectively, these islands harbor the largest number of species found in any of the geographic regions of Baja California. The number of species 1s 84, which is 48.8% of the total number (172) known from the entire peninsula and its adjacent islands. The herpetofauna of these islands is only one of two regional herpetofaunas in which the number of peninsular endemics exceeds that of the non-endemics. The other such region is the SLLR, in which the number of peninsular endemics (21) is greater than the number of non-endemics (18). In the Gulf region, the number of peninsular endemics is 1.6 times that of the non- endemics (50 vs. 32). The 50 peninsular endemics in the Gulf region include species that occupy from one to nine of the 10 recognized regions in the peninsula, as follows: Table 19. Number of herpetofaunal species in the three EVS categories (low, medium, and high) among the 10 geographic regions of the Baja California Peninsula, Mexico. Rank determined by the relative number of high EVS species. Non-native and marine Species are excluded. Geographic region CR Amphib. Reptile Conserv. November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. No. 17. Petrosaurus repens (Van Denburgh, 1895). The Short- nosed Rock Lizard ranges from Mesa San Carlos near the middle of Baja California southward to Arroyo Seco in the Isthmus of La Paz. This individual came from San Miguel Comondu, Baja California Sur. In this study its EVS was calculated as 14, placing it at the lower limit of the high vulnerability category. The IUCN determined its conservation status as Least Concern (LC), and it is not listed by SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. Lizard ranges “west of the Sierra Nevada crest from Shasta County, California, south through all of southern California west of the Mojave and Sonoran deserts” (Grismer 2002: 151) and the extreme northwestern portion of Baja California. This individual was found on the road to Sierra Juarez, Baja California, in the municipality of Ensenada. In this study its EVS was calculated as 12, placing it in the middle of the medium vulnerability category. The IUCN evaluated its conservation status as Least Concern (LC), and it is not listed by SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. Amphib. Reptile Conserv. No. 18. Petrosaurus thalassinus (Cope, 1863). The San Lucan Banded Rock Lizard “is restricted to the Cape Region of Baja California, where it occurs in at least four disjunct populations: one in the Sierra La Laguna and contiguous ranges, another in the Sierra La Trinidad, and one each on the Gulf islands of Espiritu Santo and Partida Sur...” (Grismer 2002: 149). This individual was located at Sierra Las Cacachilas, Baja California Sur, in the municipality of La Paz. In this study its EVS was assessed as 17, placing it in the middle of the high vulnerability category. The IUCN evaluated its conservation status as Least Concern (LC), and SEMARNAT determined it to be a species of Special Protection (Pr). Photo by Alan Harper. No. 20. Phrynosoma coronatum (Blainville, 1835). The Coast Horned Lizard ranges across the Cape Region of Baja California Sur and across the Magdalena Plain to the southern edge of the Vizcaino Desert (Leaché et al. 2009). This individual was found in the Sierra Las Cacachilas, Baja California Sur, in the municipality of La Paz. In this study its EVS was assessed as 12, placing it in the upper portion of the medium vulnerability category. The IUCN judged its conservation status as Least Concern (LC), and it is not listed by SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula one (38 species, 76.0%); three (two species, 4.0%); four (two species, 4.0%); five (three species, 6.0%); seven (one species, 2.0%); eight (three species, 6.0%); and nine (one species, 2.0%). Based on the relative number of high vulnerability species (Table 19), the 1‘trank 1s held by the GIR (45 of 76 species, 59.2%), the same rank as for the relative number of country endemics (see above). The rankings are the same for seven of the 10 regions. The high vulnerability ranks for species in the remaining seven regions are as follows: ATR (2™, 16 of 53 species; 30.2%); CGCR (3%, 15 of 54 species; 27.8%); SLLR (4", 13 of 39 species; 33.3%); VR (5", 10 of 58 species; 17.2%); PIR (6", 10 of 41 species; 24.4%): MR (7", nine of 45 species; 20.0%); LCVR (8", six of 59 species; 10.2%); CR (9", five of 53 species; 9.4%); and BCCFR (10", four of 27 species; 14.8%). Based on the relative numbers of peninsular endemic and high vulnerability species, the 1* rank is held by the GIR, the Gulf Islands Region, in which there are 50 peninsular endemics and 45 high vulnerability species. The peninsular endemics include 33 lizards and 17 snakes. These species are indicated with a double asterisk (see Table 4). The GIR also supports 45 high vulnerability species, which are listed below (with the EVS score in parentheses): Crotaphytus insularis** (16) Coleonyx gypsicolus** (16) Ctenosaura hemilopha** (16) Dipsosaurus catalinensis** (17) Sauromalus hispidus** (14) Sauromalus klauberi** (17) Sauromalus slevini** (16) Petrosaurus repens** (14) Petrosaurus slevini** (16) Petrosaurus thalassinus** (17) Sceloporus angustus** (16) Sceloporus grandaevus** (17) Sceloporus hunsakeri** (14) Sceloporus lineatulus** (17) Uta encantadae** (17) Uta lowei** (17) Uta squamata** (17) Uta tumidarostra** (17) Phyllodactylus bugastrolepis** (17) Phyllodactylus partidus** (16) Phyllodactylus unctus** (15) Aspidoscelis canus** (17) Aspidoscelis carmenensis** (17) Aspidoscelis catalinensis** (17) Aspidoscelis celeripes** (16) Aspidoscelis ceralbensis** (17) Aspidoscelis danheimae** (17) Aspidoscelis espiritensis** (16) Aspidoscelis franciscensis** (17) Aspidoscelis maximus** (14) Amphib. Reptile Conserv. 112 Aspidoscelis pictus** (17) Lampropeltis catalinensis** (18) Masticophis barbouri** (17) Rhinocheilus etheridgei** (17) Sonora punctatissima** (15) Sonora savagei** (16) Hypsiglena catalinae** (16) Hypsiglena gularis** (16) Hypsiglena marcosensis** (16) Crotalus angelensis** (18) Crotalus catalinensis** (19) Crotalus lorenzoensis** (19) Crotalus mitchellii (15) Crotalus polisi** (19) Crotalus thalassoporus** (19) All but one of these 45 species are peninsular endemics, and one is an MXUS species. As a group, their EVS values range from 14 to 19. The 2™ rank, with respect to high vulnerability species, is held by the ATR, the Arid Tropical Region, which includes the following 16 high vulnerability species: Elgaria paucicarinata** (14) Elgaria velazquezi** (15) Bipes biporus** (19) Ctenosaura hemilopha** (16) Petrosaurus repens** (14) Petrosaurus thalassinus** (17) Sceloporus hunsakeri** (14) Phyllodactylus unctus** (15) Phyllodactylus xanti** (15) Aspidoscelis maximus** (14) Masticophis aurigulus** (15) Sonora straminea** (15) Rena boettgeri** (14) Crotalus mitchellii** (15) Trachemys nebulosa** (15) Gopherus morafkai (15) All of these species are peninsular endemics except for one MXUS species. As a group, their EVS values range from 14 to 19. The 3™ rank is held by the CGCR, the Central Gulf Coast Region, which includes the following 15 species: Elgaria paucicarinata** (14) Elgaria velazquezi** (15) Bipes biporus** (19) Ctenosaura hemilopha** (16) Petrosaurus repens** (14) Petrosaurus thalassinus** (17) Sceloporus hunsakeri** (14) Phyllodactylus unctus** (15) Phyllodactylus xanti** (15) Aspidoscelis maximus** (14) Masticophis aurigulus** (15) November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. & : 4 oe * ie wal A Fy , ay : . A” gh i ee, ly Ms No. 21. Sceloporus angustus (Dickerson, 1919). The Isla Santa Cruz Spiny Lizard “is known only from Islas San Diego and Santa Cruz in the Gulf of California” (Grismer 2002: 158). This individual was found on Isla Santa Cruz, Gulf of California, in the municipality of Loreto. In this study its EVS is indicated as 16, placing it in the middle portion of the high vulnerability category. The IUCN judged its conservation status as Least Concern (LC), and SEMARNAT indicated it as Threatened (A). Photo by Alan Harper. No. 23. Sceloporus grandaevus (male). This individual was from Isla Cerralvo, Baja California Sur. Please see the legend for No. 22 for information on its distribution and conservation status. Photo by Tim Warfel. Amphib. Reptile Conserv. 7 No. 22. Sceloporus grandaevus (female). The Isla Cerralvo Spiny Lizard “is endemic to Isla Cerralvo in the Gulf of California in the municipality of La Paz” (Grismer 2002: 159). In this study its EVS was calculated as 17, placing in the middle portion of the high vulnerability category. The IUCN determined its conservation status as Least Concern (LC), and SEMARNAT listed it as Threatened (A). Photo by Jorge H. Valdez-Villavicencio. 4 rf OT DERLAM \ No. 24. Sceloporus licki Van Denburgh, 1895. The Cape Spiny Lizard “ranges along the mountainous foothill areas of the Cape Region from Rancho Ancon south to near La Soledad in the Sierra La Laguna...” (Grismer 2002: 173). This individual was found at Cafion San Dionisio, Sierra La Laguna, Baja California Sur, in the municipality of Los Cabos. Its EVS was determined as 13, placing it at the upper limit of the medium vulnerability category. The IUCN evaluated its conservation status as Least Concern (LC), and SEMARNAT determined it to be a species of Special Protection (Pr). Photo by Jorge H. Valdez-Villavicencio. November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Sonora straminea** (15) Rena boettgeri** (14) Crotalus mitchellii** (15) Trachemys nebulosa** (15) Gopherus morafkai (15) Of these 15 species, 14 (93.3%) are peninsular endemics, and one is a non-endemic. As a group, their EVS values range from 14 to 19. The 4" rank is held by the SLLR, the Sierra La Laguna Region, which includes the following 13 high vulnerability species: Elgaria paucicarinata** (14) Ctenosaura hemilopha** (16) Petrosaurus thalassinus** (17) Sceloporus hunsakeri** (14) Phyllodactylus unctus** (15) Phyllodactylus xanti** (15) Aspidoscelis maximum ** (14) Xantusia gilberti** (15) Masticophis aurigulus** (15) Sonora straminea** (15) Rena boettgeri** (14) Crotalus mitchellii** (15) Trachemys nebulosa** (15) All of these species are peninsular endemics. As a group, their EVS values range from 14 to 17. The 5" rank is held by the VR, the Vizcaino Region, which includes the following 10 high vulnerability species: Elgaria velazquezi** (15) Anniella geronimensis** (14) Bipes biporus** (19) Petrosaurus repens** (14) Phrynosoma cerroense** (14) Urosaurus lahtelai** (16) Aspidoscelis labialis** (15) Arizona pacata** (14) Crotalus mitchelli** (15) Trachemys nebulosa** (15) Of these 10 species, all are are peninsular endemics (76.9%), and three are non-endemics. As a group, their EVS values range from 14 to 19. The 6" rank is held by the PIR, the Pacific Islands Regions, which includes the following 10 high vulnerability species: Aneides lugubris (14) Batrachoseps major (14) Elgaria cedrosensis** (16) Amphib. Reptile Conserv. Elgaria nana** (16) Anniella geronimensis** (14) Bipes biporus** (19) Phrynosoma cerroense** (14) Lampropeltis herrerae** (20) Pituophis insulanus** (17) Crotalus mitchellii** (15) Of the 10 species in the PIR, eight are peninsular endemics and two are non-endemics. As a group, their EVS values range from 14 to 20. The 7" rank is held by the MR, the Magdalena Region, which includes the following nine high vulnerability species: Elgaria velazquezi** (14) Bipes biporus** (14) Ctenosaura hemilopha** (18) Petrosaurus repens** (14) Phrynosoma cerroense** (16) Xantusia sherbrookei** (16) Arizona pacata** (14) Crotalus mitchellii** (15) Trachemys nebulosa** (18) All nine of these species are peninsular endemics. As a group, their EVS values range from 14 to 18. The 8" rank is held by the LCVR, the Lower California Valley Region, which includes the following six high vulnerability species: Crotaphytus grismeri** (16) Phrynosoma mceallii (15) Uma notata (15) Urosaurus graciosus (15) Sonora annulata (14) Crotalus cerastes (16) Of these six species, five are non-endemics and one 1s a peninsular endemic. As a group, their EVS values range from 14 to 16. The 9" rank is held by the CR, the California Region, which includes the following five high vulnerability species: Aneides lugubris (14) Batrachoseps major (14) Anniella geronimensis** (14) Phrynosoma cerroense** (14) Aspidoscelis labialis** (15) Three of these species are peninsular endemics, and the other two are non-endemics. As a group, their EVS values range from 14 to 15. November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. No. 25. Sceloporus vandenburgianus Cope, 1896. The Southern Sagebrush Lizard occurs “from the Coast Ranges in Los Angeles County, California, south to southern San Diego County. A disjunct population occurs in the Sierra Juarez and Sierra San Pedro Martir, Baja California, Mexico...” (Grismer 2002: 174). This individual came from La Tasajera, Sierra San Pedro Martir, in the municipality of Ensenada. In this study its EVS is noted as 14, placing it at the lower limit of the high vulnerability category. The IUCN judged its conservation status as Least Concern (LC), and SEMARNAT assessed it as a species of Special Protection (Pr). Photo by Jorge H. Valdez- Villavicencio. Leaf-toed Gecko “on the peninsula...is restricted to the Cape Region. In the Gulf of California, it has been reported from Islas Partida Sur, Espiritu Santo, Ballena, Gallo, Gallina, and Cerralvo...” Grismer 2002: 209). This individual was found in the Sierra Las Cacachilas, Baja California Sur, in the municipality of La Paz. In this study its EVS is indicated as 15, placing it in the lower portion of the high vulnerability category. The IUCN judged its conservation status as Near Threatened (NT), and SEMARNAT assessed this gecko as a species of Special Protection (Pr). Photo by Jorge H. Valdez- Villavicencio. Amphib. Reptile Conserv. No. 26. Urosaurus nigricaudus (Cope, 1864). The Black- tailed Brush Lizard “ranges along the eastern side of the Peninsular Ranges from San Diego County, California, south to the Cape Region of Baja California Sur” (Grismer 2002: 180). This individual was found in the Cafion San Dionisio, Sierra La Laguna, Baja California Sur, in the municipality of Los Cabos. In this study its EVS was estimated as 7, placing it in the upper portion of the low vulnerability category. The IUCN judged its conservation status at Least Concern (LC), and SEMARNAT listed it as Threatened (A). Photo by Jorge H. Valdez-Villavicencio. ai ray - ‘ ‘ *; : img ; | ; BE Es awe i ie Pere WRB Se s i No. 28. Plestiodon lagunensis Van Denburgh, 1895. The San Lucan Skink “has a disjunct distribution throughout southern Baja California...In the Cape Region, it is restricted to the Sierra La Laguna and associated eastern foothills. North of the Cape Region, it is known from four localities: the vicinity of the Comondts; Santa Agueda, approximately 150 km to the north...; northern Sierra Guadalupe; and San Francisco de la Sierra...” (Grismer 2002: 237-238). This individual came from La Purisima, Baja California Sur, in the municipality of Comondu. In this study its EVS was judged as 13, placing it at the upper limit of the medium vulnerability category. The IUCN assessed its conservation status as Least Concern (LC), and SEMARNAT indicated it as a species of Special Protection (Pr). Photo by Jorge H. Valdez-Villavicencio. November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula The 10" rank is held by the BCCFR, the Baja California Coniferous Forest Region, which includes the following four high vulnerability species: Batrachoseps major (14) Sceloporus vandenburgianus (14) Lampropeltis multifasciata (14) Thamnophis elegans (14) All four of these species are non-endemics, and they all have EVS values of 14. Natural Protected Areas in the Baja California Peninsula and its Adjacent Islands Natural Protected Areas In Mexico, natural protected areas (ANPs, from the Spanish acronym for “Areas Naturales Protegidas” or NPAs for “Natural Protected Areas” in English) are those established to preserve natural environments among the different landscapes of the country for the conservation of endemic and endangered species, as well as for maintaining their genetic diversity and for promoting sustainable use and scientific research. The Baja California Peninsula contains 30 NPAs, which are divided into the following six categories: National Parks, Biosphere Reserves, Flora and Fauna Protection Areas, State Parks, State Reserves, and Voluntarily Designated Conservation Areas (Areas Designadas Voluntariamente ala Conservacion, or ADVC for the Spanish acronym). Of these, 17 are administrated at the federal level and two at the state level, while the remaining 11 are private (Table 20). Twenty-five of the NPAs are terrestrial and five are marine reserves. The NPAs were created between the years 1947 and 2022, with Sierra San Pedro Martir National Park being the first reserve established in the Baja California Peninsula. Terrestrial NPAs cover slightly more than 73,000 km’, which corresponds to 49.5% of the total area of the Baja California Peninsula and its associated islands. The marine protected areas cover about 17,268 km? of marine habitat. Two NPAs are the largest in Mexico. The El Vizcaino Biosphere Reserve in Baja California Sur is the largest in the country, followed by the Area de Proteccion de Flora y Fauna Valle de los Cirios in the state of Baja California, and each of these NPAs covers more than 20,000 km/? of areal extension. Together they cover 56% of the total area designed for the protection of biodiversity and natural resources in the peninsula. In addition to the considerable protected land area represented by the NPAs, these areas are distributed in all 10 of the physiographic regions throughout the peninsula (Table 20), and thus represent all of its ecosystems. Eleven natural protected areas are privately owned, primarily in such large areas as Valle de los Cirios and El Vizcaino, which contain several towns and Amphib. Reptile Conserv. ranches. Although conservation objectives often are not compatible with the activities of the landowners (e.g., agriculture and cattle ranching), the federal governmental institutions promote agreements with them on the proper use of their resources, as well as in environmental education to properly carry out the management programs. Some of the Pacific Islands are inhabited, such as Cedros, Natividad, Guadalupe, Magdalena, and Santa Margarita, but the users do not own the land; and in most cases their use is primarily dedicated to fishing activities. For this reason, several NGOs collaborate with the staff of the NPAs to carry out adequate management programs and environmental education for the proper management of the resources, and also to avoid the introduction of non-native species into the islands. In areas such as the Sierra San Pedro Martir, however, relationships with the landowners are complicated, as they use the core areas of the NPAs for cattle ranching. Regarding herpetofaunal inventories, only six of the 23 terrestrial areas (26.1%) have conducted them. This situation highlights the general lack of sufficient herpetological surveys in these areas. In all seven of the marine reserves, five species of turtles are known to inhabit these areas (Chelonia mydas, Caretta caretta, Lepidochelys_ olivacea, Eretmochelys imbricata, and Dermochelys coriacea), in addition to the Yellow- bellied Sea Snake (Hydrophis platurus). Only one of these marine areas, Complejo Lagunar Ojo de Liebre, is located in the Pacific Ocean, and the remainder are in the Gulf of California. Effectiveness of the Natural Protected Areas in the Baja California Peninsula and Its Adjacent Islands The available information on the distribution of the herpetofaunal species known to occur within the NPAs in the Baja California Peninsula and its adjacent islands are shown in Table 21, and the results are summarized in Table 22. The data in these tables indicate that 157 (91.3%) of the 172 total species known from this region of Mexico have been recorded in one to 25 of the 30 NPAs. Thus, 15 species recorded from the Baja California Peninsula and its adjacent islands have not been recorded from any of the NPAs, and therefore are not included in Table 21. These 15 species are: Anaxyrus woodhousii, Incilius alvarius, Smilisca baudinii***, Xenopus laevis***, Lithobates berlandieri***, L. forreri***, L. yavapaiensis, Crotaphytus grismeri**, Gehyra mutilata***, Uma _ notata, Xantusia sherbrookei**, Thamnophis marcianus, Kinosternon integrum*** Gopherus morafkai, and Apalone spinifera***, Seven of these 15 species (indicated by triple asterisks) are non-native to the Baja California Peninsula and its adjacent islands, and thus are not desirable for inclusion in any of the NPAs. Therefore, only eight species (the two country endemics indicated by double asterisks and the six non-endemics) should November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. No. 29. Aspidoscelis catalinensis. The Isla Santa Catalina Whiptail “is endemic to Isla Santa Catalina, in the Gulf of California, Baja California Sur” (Grismer 2002: 230). In this study its EVS was estimated as 17, placing it in the middle portion of the high vulnerability category. The IUCN judged its conservation status as Vulnerable (VU), and SEMARNAT assessed it as a species of Special Protection (Pr). Photo by Tim Warfel. No. 31. Aspidoscelis labialis (Stejneger, 1890). The Baja California Whiptail “ranges along a narrow strip of the Pacific coast, from Punta San José just south of Ensenada south to at least 6 km southeast of Guerrero Negro. It generally extends no more than 16 km inland...in the northern Vizcaino Desert and even less so farther north” (Grismer 2002: 220). This individual was found at Guerrero Negro, Baja California Sur, in the municipality of Mulegé. In this study its EVS was estimated as 15, placing it in the lower portion of the high vulnerability category. The IUCN judged its conservation status as Vulnerable (VU), and SEMARNAT listed it as a species of Special Protection (Pr). Photo by Jorge H. Valdez-Villavicencio. Amphib. Reptile Conserv. throated Whiptail occurs “from Orange and San Bernardino counties of southern California south to Cabo San Lucas...” (Grismer 2002: 212). This individual came from Rancho Meling, Sierra San Pedro Martir, Baja California, in the municipality of San Quintin. In this study its EVS was determined as 10, placing it at the lower limit of the medium vulnerability category. The IUCN evaluated its conservation status as Least Concern (LC), and SEMARNAT judged it as Threatened (A). Photo by Jorge H. Valdez-Villavicencio. No. 32. Aspidoscelis maximus (Cope, 1863). The Cape Region Whiptail is distributed in the Cape Region from Bahia La Paz to Cabo San Lucas, including the Gulf islands of Partida Sur and Espiritu Santo (Grismer 2002). This individual was located at Sierra Las Cacachilas, in the municipality of La Paz. In this study its EVS was calculated as 14, placing it at the lower limit of the high vulnerability category. The IUCN has not assessed the conservation status of this species, but it was evaluated by SEMARNAT as a species of Special Protection (Pr). Photo by Alan Harper. 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Reptile Conserv. The herpetofauna of the Baja California Peninsula No. 33. Xantusia henshawi Stejneger, 1893. The Granite Night Lizard “ranges from western Riverside County, California, south into northwestern Baja California, to at least Cafion el Cajon of the Sierra San Pedro Martir in the east and Valle la Trinidad in the west...” (Grismer 2002: 233). This individual was found at Rancho Meling, Sierra San Pedro Martir, in the municipality of San Quintin. Its EVS was indicated as 11, placing it in the lower portion of the medium vulnerability category. The IUCN determined its conservation status as Least Concern (LC), but this species is not listed by SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. No. 35. Lichanura trivirgata Cope, 1861. The Rosy Boa “ranges widely throughout the Mojave and Sonoran deserts of the southwestern United States and northern Mexico, as well as coastal regions of southern California...In Baja California, L. trivirgata occurs in all areas...except for the upper eleva- tions of the northern Peninsular Ranges...and the Sierra La Laguna” (Grismer 2002: 260). This individual was found on Isla Cedros, in the municipality of Ensenada. In this study its EVS was estimated as 10, placing it at the lower limit of the medium vulnerability category. The IUCN judged its conservation status as Least Concern (LC), and SEMARNAT calculated it as Threatened (A). Photo by Jorge H. Valdez- Villavicencio. Amphib. Reptile Conserv. 120 No. 34. Xantusia wigginsi Savage, 1952. Wiggins’ Night Lizard ranges from southern San Diego County, California southward to northeastern Baja California Sur (http://californiaherps. com; accessed 11 December 2022). This individual came from south of Catavifia, Baja California, in the municipality of San Quintin. The species EVS was determined as 11, placing it at the lower portion of the medium vulnerability category. This species is not listed by either the IUCN or SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. No. 36. Arizona pacata Klauber, 1946. The Peninsular Glossy Snake “ranges along the Pacific coast of the southern two-thirds of Baja California from at least the turnoff to Bahia de los Angeles south to 20 km north of La Paz” (Grismer 2002: 265). This individual was found in Guerrero Negro, Baja California Sur, in the municipality of Mulegé. In this study its EVS was determined as 14, placing it at the lower limit of the high vulnerability category. The IUCN assessed its conservation status as Least Concern (LC), but this snake is not listed by SEMARNAT. Photo by Tim Warfel. ae td November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Shee’ Ee aes No. 37. Bogertophis rosaliae (Mocquard, 1899). The Baja California Rat Snake occurs “throughout the rocky slopes of the Peninsular Ranges from Mountain Springs, San Diego County, California, south to Cabo San Lucas...” (Grismer 2002: 266). This individual came from the Sierra Las Cacachilas, Baja California Sur, in the municipality of La Paz. In this study its EVS was estimated as 11, placing it in the medium vulnerabilty category. The IUCN assessed its conservation status as Least Concern (LC), but this species is not listed by SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. ‘Gee ee lus (Cope, 1861). The Cape Striped Racer “is known only from the Cape Region of Baja California, along the eastern slopes of the Sierra La Laguna...” (Grismer 2002: 286). This individual came from San Bartolo, Baja California Sur, in the municipality of La Paz. In this study its EVS was calculated as 15, placing it in the lower portion of the high vulnerability category. The IUCN assessed its conservation status as Least Concern (LC), and SEMARNAT determined its conservation status as Threatened (A). Photo by Jorge H. Valdez-Villavicencio. No. 39. Masticophis fuliginosus (Cope, 1895). The Baja California Coachwhip “ranges from extreme southwestern California south to Cabo San Lucas” (Grismer 2002: 290). This individual was encountered in the Sierra Las Cacachilas, Baja California Sur, in the municipality of La Paz. In this study its EVS was estimated as 9, placing it at the upper limit of the low vulnerability category. This species is not listed by either the IUCN or SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. Amphib. Reptile Conserv. 121 No. 40. Pituophis vertebralis (Blainville, 1835). The Baja California Gopher Snake “is endemic to Baja California peninsula, ranging continuously throughout cismontane areas from at least 43 km south (by road) of El Rosario south to Cabo San Lucas...” (Grismer 2002: 298). This individual was found in the Sierra Las Cacachilas, Baja California Sur, in the municipality of La Paz. In this study its EVS was determined as 13, placing it at the upper limit of the medium vulnerability category. The IUCN assessed its conservation status as Least Concern (LC), but this snake is not listed by SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 21. Distribution of herpetofaunal species in the Natural Protected Areas of the Baja California Peninsula, Mexico. Abbreviations are as follows: * = species endemic to Mexico; ** = species endemic to Baja California; and *** = non-native species. The numbers of the Natural Protected Areas signify the following: 1 = Constitucion de 1857; 2 = Alto Golfo y Delta del Rio Colorado; 3 = Arroyo San Miguel; 4 = Sierra San Pedro Martir; 5 = Los Montes de San Pedro; 6 = Reserva Natural San Quintin; 7 = Reserva Natural Punta Mazo; 8 = Reserva Natural Monte Ceniza; 9 = Reserva Natural Valle Tranquilo; 10 = Valle de los Cirios; 11 = Costa Salvaje Wildlands; 12 = Islas del Golfo de California; 13 = Isla Guadalupe; 14 = Islas del Pacifico de la Peninsula de Baja California; 15 = Zona Marina del Archipielago de San Lorenzo; 16 = Zona Marina Bahia de los Angeles, Canales de Ballenas y Salsipuedes; 17 = Complejo Lagunar Ojo de Liebre; 18 = El Vizcaino; 19 = Servidumbre Ecologica Rancho San Cristébal-Majiben; 20 = Bahia de Loreto; 21 = Reserva Ecologica Llanos de Magdalena; 22 = Reserva Natural El Portezuelo; 23 = Santa Martha; 24 = Zona marina del Archipiélago de Espiritu Santo; 25 = Balandra; 26 = Non- Such; 27 = Sierra La Laguna; 28 = Cabo Pulmo; 29 = Estero de San José del Cabo; and 30 = Cabo San Lucas. Te PP Poof Pos [as [as [6] [8 [9 [a0 [aa [22 [a3 [2a 25 [20 [27 [28 09 pemmncmss TET PETE Anura (11 species) ANAXYVUus boreas = Bufonidae (4 species) [| fal Anaxyrus californicus Anaxyrus cognatus + Anaxyrus punctatus al [id A ylidae (2 species) [| Ea eel Fa] Pseudacris cadaverina Pseudacris hypochondriaca Ranidae (3 species) ithobates catesbeianus*** ie a Rana draytonii Rana boylii Scaphiopodidae (2 species) caphiopus couchii + ii + Caudata (3 species) Plethodontidae (3 species) im a a - Ea LJ f) Aneides lugubris | 3) al = | [| [al a Batrachoseps major nsatina eschscholtzii REPTILIA (143 species) Squamata (135 species) Anguidae (5 species) Igaria cedrosensis** Igaria multicarinata + Igaria nana** Igaria paucicarinata** Igaria velazquezi* A nniellidae (2 species) Anniella geronimensis** Anniella stebbinsi Bipedidae (1 species) Bipes biporus** T Crotaphytidae (4 species) Crotaphytus insularis** Crotaphytus vestigium Gambelia copeii** Cl [al [EJ] [ial [ [ial ia fl ii] a iS i fl Ea i [=| = i [= ] [=] [mal bz] Gambelia wisli Eublepharidae (3 species) [| Coleom gypsicous** || P(e aC Li Coleonx switak fale ca isa] hes PS ST esi ft tS] aS et EP aed Pe PS Pa a Zenit pea hammondii Amphib. Reptile Conserv. 122 November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Lt ee aki Bre cs ee No. 41. Salvadora hexalepis (Cope, 1866). The Western Patch- nosed Snake is distributed “through much of the American southwest and most of northwestern Mexico. In Baja California, S. hexalepis ranges throughout the entire peninsula. ..except for the upper elevations of the Sierra Juarez and Sierra San Pedro Martir...It is known from the Pacific islands of San Geronimo and Todos Santos and from the Gulf islands of Espiritu Santo, San José, and Tiburon” (Grismer 2002: 303). This individual was found in the Sierra Las Cacachilas, Baja California Sur, in the municipality of La Paz. Its EVS was calculated as 9, placing it at the higher limit of the low vulnerability category. The IUCN assessed its conservation status as Least Concern (LC), but this species is not listed by SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. § No. 43. Sonora fasciata (Cope, 1892). The Banded Sand Snake is distributed “from the central Baja California Peninsula at least as far north as Las Tres Virgenes to southern Baja California Sur near El Triunfo” (Cox et al. 2018). This individual came from near Loreto, Baja California Sur, in the municipality of Loreto. In this study its EVS was assessed as 12, placing it in the higher portion of the middle vulnerability category. This Species is not listed by the IUCN, but SEMARNAT lists it as a species of Special Protection (Pr). Photo by Jorge H. Valdez- Villavicencio. Amphib. Reptile Conserv. e 5 » rhe No. 42. Sonora annulata (Baird, 1859). The Shovel-nosed Snake is distributed “in the Colorado and Sonoran Deserts of Arizona and California in the USA, and in the states of Baja California and Sonora in Mexico” (Cox et al. 2018). “In Baja California, S. annulata is restricted to the Lower Colorado Valley Region, extending 34 km south of San Felipe” (Grismer 2002: 271). This individual was located at Restaurant El Michoacan, El Chinero, Baja California in the municipality of Mexicali. The species EVS was calculated as 14, placing it at the lower limit of the high vulnerability category. This snake is not listed by either IUCN or SEMARNAT. Photo by Tim Warfel. RSG No. 44. Hypsiglena slevini Tanner, 1943. The Slevin’s Night Snake occurs from near Puertecitos in the north (Murray et al. 2015) and “ranges continuously from at least Bahia de los Angeles in the north to Cabo San Lucas in the south...It is also known from Isla Santa Margarita, of the west coast of Baja California...and from Islas Cerralvo and Danzante in the Gulf of California” (Grismer 2002: 273). This individual came from Sierra la Gata, Baja California Sur, in the municipality of La Paz. In this study its EVS is noted as 11, placing it in the lower portion of the medium vulnerability category. The IUCN assessed its conservation status as Least Concern (LC), but SEMARNAT judged its conservation status as Threatened (A). Photo by Jorge H. Valdez-Villavicencio. Bs ve November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 21 (continued). Distribution of herpetofaunal species in the Natural Protected Areas of the Baja California Peninsula, Mexico. Abbreviations are as follows: * = species endemic to Mexico; ** = species endemic to Baja California; and *** = non-native species. The numbers of the Natural Protected Areas signify the following: 1 = Constitucion de 1857; 2 = Alto Golfo y Delta del Rio Colorado; 3 = Arroyo San Miguel; 4 = Sierra San Pedro Martir; 5 = Los Montes de San Pedro; 6 = Reserva Natural San Quintin; 7 = Reserva Natural Punta Mazo; 8 = Reserva Natural Monte Ceniza; 9 = Reserva Natural Valle Tranquilo; 10 = Valle de los Cirios; 11 = Costa Salvaje Wildlands; 12 = Islas del Golfo de California; 13 = Isla Guadalupe; 14 = Islas del Pacifico de la Peninsula de Baja California; 15 = Zona Marina del Archipielago de San Lorenzo; 16 = Zona Marina Bahia de los Angeles, Canales de Ballenas y Salsipuedes; 17 = Complejo Lagunar Ojo de Liebre; 18 = El Vizcaino; 19 = Servidumbre Ecologica Rancho San Cristébal-Majiben; 20 = Bahia de Loreto; 21 = Reserva Ecologica Llanos de Magdalena; 22 = Reserva Natural El Portezuelo; 23 = Santa Martha; 24 = Zona marina del Archipiélago de Espiritu Santo; 25 = Balandra; 26 = Non-Such; 27 = Sierra La Laguna; 28 = Cabo Pulmo; 29 = Estero de San José del Cabo; and 30 = Cabo San Lucas. BE [oo Po Ps [is [as [a6 [or] ow [9 [0 [aa [a2 25 [20 [8 [26 [27 [28 [29 Femina [| femidcoas ion [| fguniase Oeste) |_| Ctenoscra hemiopna®™ |_| Dinsscurasctatnensoe> |_| Dipsscurasdosats |_| sana riotopar™™ |_| Scuomaiucater |_| Scuromaius spin |_| Scuromalustanber®™ |_| a ia ia Bi a! ii = Bl al + Phrynosomatidae (29 species : Pinynosomacermense™™ | Pinyosome corona | Pinynosomamecati____| Pinynosomea planes |_| Seeopornsangnsns™= |_| Seeloporns grandaen™™ |_| Seetopornsnsaker™ |__| Seeoporwstich®™ |_| Seeopornstneantae™ |_| Seeopornsmagiver | _ Seetoporn oceidenats | = Seetopornsoreui [> Seeoporn vandenburgtms | = | Sceoporuszoxeroma*® | eli i im Fal eal al Pai Ez] i J Cl ail + BRRRERE BREE EERE BERR EERE BERR BERTGEE BERR +{ || i+] [+i TT +t +] | +{ | Tt i+] +i TT +] +] dT di Ea Ka Ei Ei + — et = = ES Phyllodactylidae (5 species) + ERR eRe ERE i eA kJ 3 I) = EY SP I eT fe a SERRE EeSEEREERReRARE BER BERDER ERE eee SEC ESRI FS aes eis) Es ES SSSI Sf ee CE I ATS) ee Se Sl Sis eas) EE SERESRERRERREREEREREERERREEe ESR A SERRE RRR BREE Amphib. Reptile Conserv. 124 November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. o No. 45. Rena boettgeri (Werner, 1899). The Cape Threadsnake is distributed at “low elevations (up to 300 m) in the Cape region of the Baja California Peninsula (Isthmus of La Paz to Cabo San Lucas) and on Isla Cerralvo” (Heimes 2016: 27). This individual came from La Paz, Baja California Sur, in the municipality of La Paz. In this study its EVS was estimated as 14, placing it at the lower limit of the high vulnerability category. This species is not listed by either the IUCN or SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. Western Terrestrial Garter Snake “ranges continuously from southwestern Canada, south through the western United States to central Nevada, Arizona, and New Mexico, nearly to the edges of the Mojave and Sonoran deserts. It is known from isolated populations in the Sierra Nevada and the San Bernardino Mountains of California as well as central New Mexico...In Baja California, 7’ elegans occurs in another isolated population from the Sierra San Pedro Martir” (Grismer 2002: 308). This individual came from La Grulla, Sierra San Pedro Martir, in the municipality of Ensenada. In this study its EVS was calculated as 14, placing it at the lower limit of the high vulnerability category. The IUCN judged its conservation status as Least Concern (LC), but SEMARNAT assessed it as Threatened (A). Photo by Jorge H. Valdez-Villavicencio. Amphib. Reptile Conserv. ee ee -* A EOE EE a Yongkang 3 = OTRO tn WSEAS No. 46. Rena humilis (Baird and Girard, 1853). The Western Threadsnake “ranges throughout the southwestern United States and western Mexico” (Heimes 2016: 30). This individual was found in Ensenada, Baja California, in the municipality of Ensenada. In this study its EVS was determined as 8, placing it in the upper portion of the low vulnerability category. The IUCN judged its conservation status as Least Concern (LC), but this threadsnake is not listed by SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. . ca wy a : wee ae, Y. MV ee ag se; Ve NS eo Se a a No. 48. Thamnophis validus (Kennicott, 1860). The West Coast Garter Snake “has a fragmented distribution along the west coast of southwestern Mexico from southern Sonora south to Guerrero...In Baja California, it also has a fragmented distribution and is known from water systems near La Burrera along the western face of the Sierra La Laguna and the watercourses and systems associated with Santiago, Agua Caliente, and Miraflores along the eastern face of the Sierra La Laguna” (Grismer 2002: 312). This individual is from Miraflores, Baja California Sur, in the municipality of Los Cabos. In this study its EVS was determined as 11, placing it in the lower portion of the medium vulnerability category. The IUCN assessed its conservation status as Least Concern (LC), but this species is not listed by SEMARNAT. Photo by Tim Warfel. November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 21 (continued). Distribution of herpetofaunal species in the Natural Protected Areas of the Baja California Peninsula, Mexico. Abbreviations are as follows: * = species endemic to Mexico; ** = species endemic to Baja California; and *** = non-native species. The numbers of the Natural Protected Areas signify the following: 1 = Constitucion de 1857; 2 = Alto Golfo y Delta del Rio Colorado; 3 = Arroyo San Miguel; 4 = Sierra San Pedro Martir; 5 = Los Montes de San Pedro; 6 = Reserva Natural San Quintin; 7 = Reserva Natural Punta Mazo; 8 = Reserva Natural Monte Ceniza; 9 = Reserva Natural Valle Tranquilo; 10 = Valle de los Cirios; 11 = Costa Salvaje Wildlands; 12 = Islas del Golfo de California; 13 = Isla Guadalupe; 14 = Islas del Pacifico de la Peninsula de Baja California; 15 = Zona Marina del Archipielago de San Lorenzo; 16 = Zona Marina Bahia de los Angeles, Canales de Ballenas y Salsipuedes; 17 = Complejo Lagunar Ojo de Liebre; 18 = El Vizcaino; 19 = Servidumbre Ecolégica Rancho San Cristobal-Mayjiben; 20 = Bahia de Loreto; 21 = Reserva Ecologica Llanos de Magdalena; 22 = Reserva Natural El Portezuelo; 23 = Santa Martha; 24 = Zona marina del Archipiélago de Espiritu Santo; 25 = Balandra; 26 = Non- Such; 27 = Sierra La Laguna; 28 = Cabo Pulmo; 29 = Estero de San José del Cabo; and 30 = Cabo San Lucas. BET [eof Ps [as [as [ae [or] ow [09 [0 [aa [ae [25 20 [8 [26 [27 [28 [29 Pintodaannspane [| | |_| a Pitodacyus nese |_| Pitodacyussamar™ |_| Senckne Gps) |_| Peston ibers_____[ + | Peston togineniee |_| Peston stitontame —_[* | shane (Sapien |_| spose cams |_| spose cameron |_| spose catatnenis™™ |_| ial = im il = im El ia Fal Es Kantasine Gapecy |_| Fama giterine |_| Fania viegins |_| Charniine spec |_| + Lamoopeisctatnensisee |_| Lameopetihererae™® |_| + SERRE oC OE CR ee BRRERREEREEEREREEE 2 BEE BERR Be Amphib. Reptile Conserv. 126 November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. = a No. 49. Crotalus angelensis Klauber, 1963. The Isla Angel de la Guarda Rattlesnake “is endemic to Isla Angel de la Guarda in the Gulf of California” (Grismer 2002: 333), from where this individual originated. In this study its EVS was assessed as 18, placing in the upper portion of the high vulnerability category. The IUCN judged its conservation status as Least Concern (LC), but this rattlesnake is not listed by SEMARNAT. Photo by Tim Warfel. No. 51. Crotalus polisi Meik, Schaack, Flores-Villelam and Streicher, 2018. The Isla Cabeza de Caballo Speckled Rattlesnake or Horsehead Island Speckled Rattlesnake is endemic to Isla Cabeza de Caballo Island in the Gulf of California, municipality of San Quintin. The species EVS was calculated as 19. The species is not listed by the IUCN, but SEMARNAT assessed its conservation status as Special Protection (Pr). Photo by Tim Warfel. Amphib. Reptile Conserv. fae ea oy or 7 . ba a he a No. 50. Crotalus enyo (Cope, 1861). The Baja California Rattlesnake “ranges throughout most of Baja California. In the north, its contact with the Pacific coast occurs in the vicinity of Cabo Colonet and with the Gulf coast near Bahia de los Angeles. From here, C. enyo continues south throughout all of Baja California...It is also known from the Pacific islands of Magdalena and Santa Margarita and the Gulf islands of Carmen, Cerralvo, Coronados, Espiritu Santo, Pardo, Partida Sur, San Francisco, San José, and San Marcos...” (Grismer 2002: 328-329). This individual was found at Guerrero Negro, Baja California Sur, in the municipality of Mulegé. In this study its EVS was estimated as 13, placing it at the upper limit of the medium vulnerability category. The IUCN assessed its conservation status as Least Concern (LC), but SEMARNAT judged its status as Threatened (A). Photo by Jorge H. Valdez- Villavicencio. rot Aa, Lie ? - Aa thew, by aie a) ae oe No. 52. Crotalus pyrrhus (Cope, 1866). The Southwestern Speckled Rattlesnake is distributed from southeastern Nevada, western Arizona, and southern California southward into the northern half of the Baja California Peninsula (Meik et al. 2015). This individual is from Rancho La Costilla, Sierra San Pedro Martir, in the municipality of San Quintin. Its EVS was assessed as 13, placing it at the upper limit of the medium vulnerability category. This species is not listed by the IUCN, but it is listed by SEMARNAT as a species of Special Protection (Pr). Photo by Ivan Parr. November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 21 (continued). Distribution of herpetofaunal species in the Natural Protected Areas of the Baja California Peninsula, Mexico. Abbreviations are as follows: * = species endemic to Mexico; ** = species endemic to Baja California; and *** = non-native species. The numbers of the Natural Protected Areas signify the following: 1 = Constitucién de 1857; 2 = Alto Golfo y Delta del Rio Colorado; 3 = Arroyo San Miguel; 4 = Sierra San Pedro Martir; 5 = Los Montes de San Pedro; 6 = Reserva Natural San Quintin; 7 = Reserva Natural Punta Mazo; 8 = Reserva Natural Monte Ceniza; 9 = Reserva Natural Valle Tranquilo; 10 = Valle de los Cirios; 11 = Costa Salvaje Wildlands; 12 = Islas del Golfo de California; 13 = Isla Guadalupe; 14 = Islas del Pacifico de la Peninsula de Baja California; 15 = Zona Marina del Archipielago de San Lorenzo; 16 = Zona Marina Bahia de los Angeles, Canales de Ballenas y Salsipuedes; 17 = Complejo Lagunar Ojo de Liebre; 18 = El Vizcaino; 19 = Servidumbre Ecologica Rancho San Cristobal-Majiben; 20 = Bahia de Loreto; 21 = Reserva Ecoldégica Llanos de Magdalena; 22 = Reserva Natural El Portezuelo; 23 = Santa Martha; 24 = Zona marina del Archipiélago de Espiritu Santo; 25 = Balandra; 26 = Non-Such; 27 = Sierra La Laguna; 28 = Cabo Pulmo; 29 = Estero de San José del Cabo; and 30 = Cabo San Lucas. Te ef eps Poppa [vs Jas] v6] 7 fas] 9 [a0] a1 [a2 [29 [26 [25 [20 [27 [28 [>> [a0 einem [sale olaleie|sf]e] |] | suaiorneoes + [+ |_| ee ee Oe Se re ema eae) eve lel ol he) abel ele [eed aie |e Crotalusarrox | TTL TL Crotalus caratinensis** || oT oT TT tT tT ET | t+} TT TE TE ET tT ET Crotalus cerastes | t+} | tT | tT | tT tT | TE ET Crotatusenyor* | OT TT TT TT ede ed OD Ee Ee Crotalus micchettnt® TT TT | | dT TE TT et +t ee Crotaluspotisit® | oT TT TT dT TE TT T+} ET EE TE ET EE Grotiius pyri | Pee fe Pe ey ee ea estat tiers aL eh] i [ae [a | | a lee [ef tise ef | ee [ea Crotalus thatassoporus* ||} | TT oT tT tT TT t+} TE TT TE TE TE TT TE restuaines (species) | {| | | | | | | | | | |} PP tT EE EE ET ET EE Amphib. Reptile Conserv. 128 November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. i hi *" ‘ {i s< No. 53. Crotalus ruber Cope, 1892. The Red Diamond Rattlesnake is distributed from “Los Angeles County, California, south throughout Baja California to Cabo San Lucas...Crotalus ruber is known to occur on the Pacific islands of Cedros and Santa Margarita...and the Gulf islands of Angel de la Guarda, Danzante, Monserrat, Pond, San José, and San Marcos...” (Grismer 2002: 322). This individual was found at Rancho El Coyote, Sierra San Pedro Martir. The species EVS was determined as 9, placing it at the upper limit of the low vulnerability category. The IUCN assessed its conservation staus as Least Concern (LC), and SEMARNAT judged it as a species of Special Protection (Pr). Photo by Ivan Parr. No. 55. Actinemys palli da (Seeliger, 1945). The Southwestern Pond Turtle ranges from the central coast range south of the San No. 54. Crotalus thalassoporus Meik, Schaack, Flores-Villela, and Streicher, 2018. The Piojo Island Speckled Rattlesnake or Louse Island Speckled Rattlesnake is endemic to Isla Piojo in the Gulf of California, in the municipality of San Quintin. Its EVS was assessed as 19, placing it in the upper portion of the high vulnerability category. This species is not listed by the IUCN, but SEMARNAT lists it as a species of Special Protection (Pr). Photo by Tim Warfel. a Francisco Bay area to northwestern Baja California (http://www.reptile-database.org/, accessed 7 December 2022: Grismer 2002: 88), with a disjunct population in the central desert of Baja California (Valdez-Villavicencio et al. 2016). This individual came from Arroyo Valladares, Baja California, in the municipality of San Quintin. In this study its EVS was assessed as 13, placing it at the higher limit of the medium vulnerability category. This species is not listed by either the IUCN or SEMARNAT. Photo by Jorge H. Valdez-Villavicencio. Amphib. Reptile Conserv. 129 November 2023 | Volume 17 | Number 1 & 2 | e326 The herpetofauna of the Baja California Peninsula Table 21 (continued). Distribution of herpetofaunal species in the Natural Protected Areas of the Baja California Peninsula, Mexico. Abbreviations are as follows: * = species endemic to Mexico; ** = species endemic to Baja California; and *** = non-native species. The numbers of the Natural Protected Areas signify the following: 1 = Constitucion de 1857; 2 = Alto Golfo y Delta del Rio Colorado; 3 = Arroyo San Miguel; 4 = Sierra San Pedro Martir; 5 = Los Montes de San Pedro; 6 = Reserva Natural San Quintin; 7 = Reserva Natural Punta Mazo; 8 = Reserva Natural Monte Ceniza; 9 = Reserva Natural Valle Tranquilo; 10 = Valle de los Cirios; 11 = Costa Salvaje Wildlands; 12 = Islas del Golfo de California; 13 = Isla Guadalupe; 14 = Islas del Pacifico de la Peninsula de Baja California; 15 = Zona Marina del Archipielago de San Lorenzo; 16 = Zona Marina Bahia de los Angeles, Canales de Ballenas y Salsipuedes; 17 = Complejo Lagunar Ojo de Liebre; 18 = El Vizcaino; 19 = Servidumbre Ecologica Rancho San Cristébal-Majiben; 20 = Bahia de Loreto; 21 = Reserva Ecoldégica Llanos de Magdalena; 22 = Reserva Natural El Portezuelo; 23 = Santa Martha; 24 = Zona marina del Archipiélago de Espiritu Santo; 25 = Balandra; 26 = Non-Such; 27 = Sierra La Laguna; 28 = Cabo Pulmo; 29 = Estero de San José del Cabo; and 30 = Cabo San Lucas. ET Poo Po Ps Ps Pas [a6 [07 [a8 [9 [0 [au [a2 [25 [2s [25 [26 [ar [28 [> [a0 See Cre ee a a fremooneisinoricna | | ||111]111/[ | Lenicchepw oivacea | | | ||| [11] ie] | Dermocrewinecrwverey | | |||) >PPt yl ermochelys coriacea | |_| | | | | | | | | t+] tet+t+t+] | te} tT tet tt dey fe Emydidae@species) | | | | | | | | | tT | | | | tT | tT | TT EE TE Herinemys marmorata || | t+ tT +t ToT | tet} | PT tT tT TE ET EE ET rachemysnebuiosa** | | | ot tT tT tT tT tT tT TE EE ET TT T+#T TE TE rachemys scripta*** || TL TT TT cE TT dP TT TT TT TE frotatcis7species) | tT | | | | PT | PT PT PTE PE TT EE TE EE EE eventually be located within one or more of the NPAs. The most widely distributed species among the NPAs is the phrynosomatid lizard Uta stansburiana, which has been recorded from 25 of these areas. A sizeable number of species (69 or 43.9%) have been recorded from only one of the 30 NPAs (Table 21). The remaining 90 species (57.3%) have been recorded in two to 24 NPAs. In 29 of the 30 NPAs from which herpetofaunal species have been recorded, the total numbers of species range from six to 84 (Table 22). The allocations of these species according to their distributional status are shown in Table 22. Most of the 29 NPAs (22 or 75.9%) lack non-native species, which is a desirable feature. The other seven NPAs support from one to four non-native species (x = 1.7 species). Each of the 29 NPAs have a herpetofauna that contains some number of non-endemic species, which ranges from six to 40. Most of these NPAs also have some country endemic species, ranging from one to 50; however, no country endemic species have been recorded in eight of these areas. Usually the number of non-endemic species exceeds that of the country endemics, except for the Gulf of California Islands (with 32 of the former and 50 of the latter), the Sierra La Laguna (with 18 of the former and 20 of the latter), and the Estero de San José del Cabo (with 15 of each of non- endemic and country endemic species). Only one of the 30 NPAs in the Baja California Peninsula and its adjacent islands (Isla Guadalupe) lacks a herpetofaunal survey of any magnitude. This island 1s located 241 km off the western coast of the peninsula and about 400 km southwest of Ensenada in the state of Amphib. Reptile Conserv. Baja California. This island and its surrounding islets and waters are part of a biosphere reserve established in 2005 (Table 20). The absence of herpetofaunal documentation on Isla Guadalupe is not due to a lack of effort on the part of herpetologists, as several collecting efforts have been made. For example, the San Diego Natural History Museum sponsored an expedition there years ago, and two authors of this paper (APG and JHVV) also have been on this island and contacted people who are part of an NGO at a base camp. However, none of these efforts resulted in the finding of any amphibians or reptiles on Isla Guadalupe; therefore, it is interesting that apparently no amphibians, and especially no reptiles, have made it to this island. Of the 81 non-endemic species found in the herpetofauna of the peninsula and its associated islands, 75 (92.6%) have been recorded among the region’s NPAs. Of the 77 country endemic species known, 76 (98.7%) are established in protected areas. Fortunately, only seven of the 14 non-native species (50%) have populations established in any of the NPAs, and usually they occur individually in any given area. Only one of these seven non-native species (Hemidactylus frenatus) occurs in more than a single NPA, and the number of areas it occurs in 1s six (Table 21). The Baja California Peninsula is a unique area of Mexico, as nearly 30% of its land is protected. However, comprehensive herpetological studies are still lacking for most reserves, and the available surveys largely focus on target species (endemics and NOM-S59 species). Since several conservation threats affect the natural protected November 2023 | Volume 17 | Number 1 & 2 | e326 Anny Peralta-Garcia et al. Table 22. Summary of the distributional status of the herpetofaunal species in protected areas in the Baja California Peninsula, Mexico, and adjacent islands. Total = total number of species recorded in a compendium of the listed protected areas. Distributional status Number of Non: Cou Non-native species endemic Endemic (NN) (NE) (CE) a SS a CO a TT a A [Reserva Natural San Quintin ———OSOSCSC~—~—S—CSC‘“<~<;7) 2000-3000 |_| >3000 Boenmeee @ * 50 kilometers Sea Caribbean $4 “83 “82 Fig. 1. (A) Map showing the known populations of the species within the Bolitoglossa subpalmata species group. (B) Map showing the localities for B. bolanosi sp. nov. and the species close to it in the paramo of the Cordillera de Talamanca. The open shapes indicate the historical data and the solid shapes indicate the localities with molecular data included herein. the subalpine rain paramo of ICA is relatively small in area, several regions remain unexplored and several species may possibly remain unnamed. To the best of our knowledge, only three species of salamanders are exclusively found in the ICA’s paramo: Bolitoglossa kamuk, B. pesrubra, and B. pygmaea. Most salamanders of the ICA are associated with the highlands, especially on the Cordillera de Talamanca, where studies have documented high species turnover between sites and along elevational gradients (Garcia- Paris et al. 2000; Wake 1987). The Bolitoglossa subpalmata Species Group (Parra-Olea et al. 2004) is an example of this pattern, containing eight species (B. bramei, B. gomezi, B. gracilis, B. kamuk, B. pesrubra, B. splendida, B. subpalmata, and B. tica) that are distributed in the highlands of Costa Rica and western Panama (Fig. 1), with high species turnover along latitudinal and elevational gradients (AmphibiaWeb 2023; Boza- Oviedo et al. 2012; Garcia-Paris et al. 2008). Although the phylogenetic relationships of the Bolitoglossa subpalmata S.G. have been relatively well-studied (Boza-Oviedo et al. 2012), there is still little information on the distribution ranges of the species in this group, especially B. pesrubra, and the area between the paramo of the Cerro Chirrip6 and the paramo of the Cerro Kamuk has not been sampled at all. During fieldwork in the subalpine rain paramo in the Cordillera de Talamanca, specifically the summits of the peaks Durika, Arbolado, Haku, and Utyum (Fig. 1), we found some specimens of the B. subpalmata S.G., filling in the distribution range for the group. Based on the molecular and morphological data reported herein, we Amphib. Reptile Conserv. describe these samples as a new species of Bolitoglossa closely related to B. kamuk and B. pesrubra of the B. subpalmata S.G. Materials and Methods Taxon sampling. In July 2013, March 2015, January 2016, and October 2016, we carried out collecting trips to Cerro Utyum (July 2013 and March 2015), Cerro Durika (January 2016), Cerro Arbolado (October 2016), and Cerro Haku (October 2016) on the Talamanca Mountain range within La Amistad International Park, Costa Rica (Fig. 1). These trips involved walking a transect from Olan, Buenos Aires (9.2788° N, 83.2156° W, 1,500 m asl; all GPS coordinates in WGS84 datum) on the Pacific slope to the summits of the peaks. The salamanders were found by opening bromeliads from previously cut-off trees, and a total of 19 specimens referred to Bolitoglossa subpalmata S.G. were found in the sampled peak summits. All the specimens collected for this study were humanely euthanized using a _ topical anesthetic. Following the euthanization process, a small tissue sample, either liver or the tip of the tail, was taken and stored in 96% ethanol. The specimens were fixed in a 10% formalin solution and transferred to 70% ethanol for long-term preservation. Individuals were deposited in the herpetological collection of Museo de Zoologia at Universidad de Costa Rica (UCR) (Appendix 1). Museum collection acronyms follow Frost (2023), with the additions of EAP (Erick Arias field numbers), CRARC (Costa Rica Amphibian Research Center private December 2023 | Volume 17 | Number 1 & 2 | e327 Arias et al. collection), and NV= No voucher. Amplification and sequencing. Total genomic DNA was extracted from the ethanol-preserved tissues of 24 Bolitoglossa specimens using the phenol-chloroform standard protocol (Sambrook and Russell 2006). The large subunit ribosomal RNA (16S) and cytochrome b (cyt 6) mitochondrial genes were amplified. The primers 16Sar and 16Sbr (Palumbi et al. 1991) were used for 16S, and primers MVZ15 and MVZ16 (Moritz et al. 1992) were used for cyt b. PCR amplifications were performed using a total volume of 15 uL, which contained 1 uL DNA template (at 50 ng uL"'), 0.75 U Taq polymerase (Amplificasa®, Biotecnologias Moleculares), 1X PCR buffer with 1.5 mM MgCl, 0.2 mM deoxynucleotide triphosphates (dNTPs), and 0.3 uM forward and reverse primers. The PCR conditions were as follows: for 16S, an initial cycle of 5 min at 94 °C, followed by 35 cycles of 45 s at 94 °C, 30 s at 55 °C, 45 s at 72 °C, plus a final step of 3 min at 72 °C; and for cyt 5, an initial cycle of 2 min at 94 °C, followed by 38 cycles of 30 s at 94 °C, 1 min at 48 °C, 1 min at 72 °C, plus a final step of 8 min at 72 °C. The PCR products were cleaned with ExoSap-IT (USB Corporation) and sequenced in both directions using the original amplification primers and BigDye termination reaction chemistry (Applied Biosystems). The cycle-sequencing products were column-purified with Sephadex G-50 (GE Healthcare) and run on an ABI 3500xL Genetic Analyzer (Applied Biosystems). Consensus sequences for each individual were constructed using SEQUENCHER 5.3 (Genes Codes Corp.). The resulting sequences were deposited in GenBank (Appendix 1). Phylogenetic analyses. The sequences obtained here were compared with the sequences of the 16S and cyt b mitochondrial genes for 78 specimens of the Bolitoglossa subpalmata species group. Sequences of B. aurae were used as outgroup and those from B. compacta were used to root all trees based on the results in Rovito et al. (2015). The list of vouchers and GenBank accession numbers used in this study are provided in Appendix 1. Sequence alignments were performed using the MUSCLE 3.7 software (Edgar 2004) with default parameters and trimmed to the point where most of the taxa had sequence data. PartitionFinder v2.1.1 software (Lanfear et al. 2017) and the Bayesian Information Criterion (BIC) were used to select the best partition scheme and the best model of sequence evolution for each partition. A single set of branchlengths was used across all partitions (branchlengths=linked), and the search for the best partition scheme used a heuristic search (scheme=greedy, Lanfear et al. 2012). Four subsets were defined a priori: one for 16S and three for cyt b (partitioned by codon position). Phylogenetic analyses were performed using both Maximum Likelihood (ML) and Bayesian Inference (BI) methods. The maximum likelihood analysis was performed using Garli 2.01 (Zwickl 2006). To find the best tree, ten search replicates were run with the following default setting values: streefname = random, attachmentsper-taxon = 24, genthreshfortopoterm = Amphib. Reptile Conserv. 100,000, and significanttopochange = 0.00001. For bootstrapping, 1,000 pseudoreplicates were run with the previous settings and with the following changes: genthreshfortopoterm = 10,000, significanttopochange =0.01, and treerejectionthreshold = 20, as suggested in the Garli manual to speed up the bootstrapping. From these bootstraps, a majority rule consensus tree was obtained using Sumtrees (Sukumaran and Holder 2010a) from DendroPy package version 4.4.0 (Sukumaran and Holder 2010b). Bayesian phylogenetic analysis was performed using MrBayes 3.2.6 (Ronquist et al. 2012) with the partition scheme and the model of sequence evolution for each partition as selected previously. Two separate analyses were run, each consisting of 50 million generations, sampled every 1,000 generations, and four chains with default heating parameters. A time-series plot of the likelihood scores of the cold chain was examined to check stationarity using Tracer 1.6 software (Rambaut et al. 2014). The first 25% of trees were discarded as burn-in and the remaining trees were used to estimate the allcompat consensus tree along with the posterior probabilities for each node and each parameter. The Garli and MrBayes analyses were run on the CIPRES portal (Miller et al. 2010). Genetic distances (uncorrected p-distances) were computed using MEGA6 (Tamura et al. 2013). Morphometrics. The measurements used herein follow those proposed by Kubicki et al. (2022). The specimens from new localities were compared morphometrically only with the species B. kamuk and B. pesrubra, given that these are the species of the group that inhabit the subalpine paramo and occur geographically close (Fig. 1, Appendix 2). The following 29 morphological measurements were taken on each of the 31 adult specimens of Bolitoglossa subpalmata S.G. that were examined in this study: standard length (SL), shoulder width (ShW), head width (HeW), neck width (NeW), eye width (EW), snout length (SnL), jaw to snout length (JSL), lateral gular fold to tip of snout (LGES), internarial distance (IND), naris to lip distance (NLP), intercanthal distance (ICD), hind limb length (HLL), front limb length (FLL), trunk width (TW), midventral gular fold to snout length (VGS), front limb to snout distance (FSL), ulna and hand length (UHL), axilla to groin length (AGL), vent length (VL), hand width (HaW), hand length (HaL), length of Finger III (LF3), width of Finger III (WF3), length of Finger II (LF2), foot width (FoW), foot length (FoL), length of Toe III (LT3), width of Toe III (WT3), and length of Toe II (LT2). The additional proportions reported here include: IND/HeW, HeW/AGL, SnL/HeW, HaL/VGS, FoL/VGS, HaW/HeW, FoW/HeW, LT2/FoL, LF2/HaL, WT3/FoW, WF3/HaW, HaL/HaW, and FoL/ Fow. The measurements were taken with a Mitutoyo Absolute 4084 digital caliper, and rounded to the nearest 0.1 mm. Given that there are very significant overlaps in the numbers of teeth found in the different Bolitoglossa species known to occur in Costa Rica (Boza-Oviedo et al. 2012; Garcia-Paris et al. 2008; Savage 2002), the number of teeth in the specimens examined in this study were not counted. We believe that attempting to accurately count December 2023 | Volume 17 | Number 1 & 2 | e327 A new species of Bolitoglossa the number of teeth in such small specimens (most of which were fixed with their mouths in a closed position), requires a high-level of invasive manipulation that would most likely cause significant and unnecessary damage to the mouth cavity. The limb interval is equal to the number of costal folds between the tips of the longest digits of the adpressed front and hind limbs, expressed 1n 0.5 increments (e.g., 4 or 4.5). Morphometric statistics. To avoid allometric effects relative to the differences in the sizes and shapes between species and between individuals, the data were transformed using the method of Lleonart et al. (2000). In this method, a logarithmic transformation of the continuous variables is performed to reduce the extreme values. All transformed variables were used in the allometric transformation by means of the equation: =n where Y* corresponds to the value of each of the dependent variables corrected for size and shape, Y corresponds to the value of each dependent morphometric variable, X, is the average of the SL variable for all populations, X, is the SL value for each individual, and 6 is the regression line intercept with the Y-axis resulting from the regression of each dependent variable with X,. The intercept is used as an allometric transformation factor and is unique for each variable. A discriminant analysis was performed to determine whether the morphometric variables were effective in predicting the species. Only those variables that were different among at least two of the three putative species analyzed were included in the discriminant analysis. The following variables were finally used in the discriminant analysis: HeW, JSL, ICD, HLL, VGS, HaW, HaL, HaL/ VGS, FoL/VGS, and HaW/HeW. The discriminant analyses were separated by sex, and were performed using R v3.3.3 (R Development Core Team 2013). Results Molecular analyses. The mitochondrial dataset includes samples of all described species in the Bolitoglossa subpalmata S.G. The resulting data matrix had a total of 80 sequences (31 salamanders collected in this study and 49 sequences downloaded from Genbank) with a sequence length of 1,334 bp, including gaps, with 527 bp for 16S and 807 for cyt b. Three partitions were identified with the following substitution models: HK Y+I+G for 16S + cyt 6 codon position 3, HK Y+I for codon position 1 of cyt b, and TRN+G for cyt 6 codon position 2. The phylogenies from Garli and MrBayes were relatively discordant in the internal topology within the B. subpalmata S.G. (Fig. 2). However, all phylogenetic analyses found the B. subpalmata S.G.to be monophyletic, with three internal clades relatively well- supported. The first clade was formed by B. bramei and B. gomezi, two species that are restricted to premontane and montane forests of the Pacific slope. The second clade was formed by B. gracilis sister to B. tica + B. Amphib. Reptile Conserv. subpalmata, and these species are distributed in the Volcanic Central Mountain range and the northern part of the Cordillera de Talamanca. Finally, a well-supported clade was formed by B. kamuk sister to B. pesrubra + unnamed taxon, and these three species are restricted to the subalpine rain paramo of the Cordillera de Talamanca. The phylogenetic position of B. splendida is uncertain. In the Bayesian analysis, this species was weakly supported as sister to the clade formed by the three species of the paramo. In the ML analysis, B. splendida was weakly supported as sister to the clade formed by B. bramei + B. gomezi, forming the sister clade to the clade formed by B. gracilis, B. subpalmata, and B. tica. The mitochondrial genetic distances are shown in Table 1. Genetic distances between the specimens of the unnamed taxon from the subalpine rain paramo and all other members of the B. subpalmata species group are 1.45—-4.71% for 16S and 5.03—9.09% for cyt b. Morphometric analyses. The specimens examined included 13 specimens (8Q and 54) of the unnamed taxon from the subalpine rain paramo, 15 specimens of B. pesrubra (7@. and 8), and three specimens of B. kamuk (18 and 23). The morphometric variation between the three species that inhabit the subalpine rain paramo of Costa Rica is shown in Table 2. The discriminant analysis correctly classified 100% of the specimens to the species (Fig. 3) in both sexes, showing a clear separation between the specimens of the new species described below and the specimens of B. kamuk and B. pesrubra. The ratios HaL/VGS, HaW/HeW, and IND/HeW differed among the three species of the subalpine rain paramo. Description of New Species Bolitoglossa bolanosi sp. nov. Bolafios’ Web-footed Salamander (Figs. 4-6) urn:Isid:zoobank.org:act:3 F7EO0D7E-906F-4FCD-B4CE-0A48F134DEA1 Holotype. UCR 22965, an adult male from Costa Rica: Provincia de Puntarenas: Canton de Buenos Aires: Distrito de Buenos Aires: the summit of Cerro Arbolado, Parque Internacional La Amistad, (9.320°, -83.216°; 2,600 m asl), collected by Erick Arias and Omar Zufiga on 19 October 2016. Paratopotype. UCR 22964, a subadult male, same data as holotype. Paratypes. UCR 22424, an adult male; UCR 22423, an adult female; UCR 22425, a subadult female; UCR 22422 and UCR 22426, subadult males; and UCR 22427, a juvenile from Costa Rica: Provincia de Puntarenas: Canton de Buenos Aires: Distrito de Buenos Aires: the summit of Cerro Utyum, Parque Internacional LaAmistad, (9.323°, -83.187°; 2,870 m asl), collected by Erick Arias, Gerardo Chaves, Olmer Cordero, and Omar Zufiga on 30 March 2015. UCR 22421, an adult female from Costa Rica: Provincia de Limon: Canton de Talamanca: Distrito de Telire: the summit of Cerro Utyum, Parque December 2023 | Volume 17 | Number 1 & 2 | e327 0.02 Arias et al. B. compacta UCR20532 B. aurae UCR22842 51 63 31 a3 96 2 N N N nn ore No Uo om) Fee ior = BODO AupApPHHp OOBAWNCO_} MVZ229170 UC R20 iB. gracilis 5 28 ae NN COE, a @ S 5 Nd 5 se NS INS hd RN aS e) G wl) OPES — Q \O B. bolanosi sp. nov. C 55 eS el SSN i) cc le Ss MAWIYeRN c wey G Lvs) Com} me (oe) DB UCR20251 NV-Cerro de la Muerte |B. pesrubra MVZ190928 MVZ225881 MVZ195598 Fig. 2. Bayesian phylogenetic inference shows the relationships of the Bolitoglossa subpalmata species group based on the 16S and cyt b mitochondrial DNA gene fragments. Bootstrap proportions from maximum likelihood are shown above the branches, and numbers below the branches are posterior probabilities (multiplied by 100) from the MrBayes analysis. The scale bar refers to the estimated substitutions per site. The support values of any node within the species are not shown. The asterisks represent support >99. NV = no voucher. Internacional La Amistad, (9.333°, -83.180°; 2,913 m asl), collected by Erick Arias, Gerardo Chaves, Olmer Cordero, and Omar Zufiiga on 30 March 2015. UCR 22745, an adult male; UCR 22741—4, adult females; and UCR 22746, a juvenile from Costa Rica: Provincia de Puntarenas: Canton de Buenos Aires: Distrito de Buenos Aires: the summit of Cerro Haku, Parque Internacional La Amistad, (9.322°, -83.203°; 2,660 m asl), collected by Erick Arias and Omar Zufiiga on 28 December 2015. Amphib. Reptile Conserv. 147 UCR 24245, an adult female; UCR 24246, an adult male; UCR 24247, a subadult male; and UCR 24248, a juvenile from Costa Rica: Provincia de Puntarenas: Canton de Buenos Aires: Distrito de Buenos Aires: the summit of Cerro Durika, Parque Internacional La Amistad, (9.374°, -83.303°; 3,240 m asl), collected by Omar Zufiiga on 13 January 2016. Generic Placement. Assigned to the genus Bolitoglossa December 2023 | Volume 17 | Number 1 & 2 | e327 A new species of Bolitoglossa due to having 14 costal grooves and lacking a sublingual Z fold, and to the subgenus Eladinea based on the molecular a evidence presented herein. g|e| 4 2 8 RBs SS auch: sab | Boy [Boel eats ceieel cg Peete kein ss whet iagnosis. The combination of the following os {8 characteristics can be used to distinguish Bolitoglossa 3 aS bolanosi from the other described species of the genus 2Is Bolitoglossa: (1) having broad hands and feet, with the iS distal phalanges on the fingers and toes free of palmar € i and plantar tissue; (2) dorsal coloration highly variable, e 3S rarely black brownish uniform and usually mottled Fe a (Wee Gretel boa tec ne | S with yellow spots, but never with red on hind limbs or Sa |beilso eee fee Pee ee te af forelimbs; and (3) 16S and cyt b mtDNA distances. oO 1s zg Comparisons. Bolitoglossa bolanosi is differentiated =) from members of the subgenus Eladinea by its 16S and eripe cyt b mtDNA distances. Since B. bolanosi is only known nD Filia & & S&S & a | 5 & to occur in Costa Rica and molecular evidence strongly ia i Oe rae ue oe te mo supports it forming part of the Bolitoglossa subpalmata s species group within the subgenus Eladinea, phenotypic 2 comparisons are presented here only concerning the S : members of that clade (B. bramei, B. gomezi, B. gracilis, Sot] tei Ore On eel at | ca Save Se B. kamutk, B. pesrubra, B. splendida, B. subpalmata, and c ax | a ~ mn a ~ Km wo B. tica), which are endemic to mountain ranges of Costa % Rica and western Panama. 8 Contrasting characteristics for Bolitoglossa bolanosi Zz x are presented in parentheses. Bolitoglossa bramei Wake S/8 et al., 2007 can be distinguished from B. bolanosi = = ao 2 ay 0 0 © & by having a rounded snout in males (snout strongly S tO: SS + STF HM HH : SLT sSl a a am A | Co KN 8 truncated in males); dorsal ground color dark brown to S| 5 brownish red, usually with darker mottling or frosting of 3 |% silvery-gray (dorsal ground color black to dark brown, = usually with mottling or blotches of yellow to red). Sle Bolitoglossa gomezi Wake et al., 2007 has fore limbs aS |e | ct -& 3 m over the soil, despite active searches in the moss of the paramo. Unlike in the peaks Durika, Arbolado, and Haku, the specimens were mainly found in the moss at <2 m over soil, despite active searches in the bromeliads at >3 m above the ground (especially on Cerro Arbolado and Cerro Haku). In the summit of Cerro Durika, B. bolanosi is very near (~4 km) to a site with B. pesrubra. In the summit of Cerro Utyum, it is to only ~7 km to a site with B. kamuk, and in this peak B. bolanosi is sympatric with an unnamed miniaturized salamander related to B. pygmaea. In addition, the type December 2023 | Volume 17 | Number 1 & 2 | e327 Arias et al. Fig. 4. Holotype of Bolitoglossa bolanosi sp. nov. (UCR 22965) in life on a white background. Photograph by Erick Arias. ie , i. ‘_f £ npr, “9 “ak ag Fig. 5. Dorsal views of the hand and foot of ae holotypes of (A-B) Bolitoglossa bolanosi sp. nov. (UCR 22965) and (C-D) B. pesrubra in preservation. Photograph by Erick Arias. Amphib. Reptile Conserv. 153 December 2023 | Volume 17 | Number 1 & 2 | e327 A new species of Bolitoglossa Fig. 6. In-life photographs of variations within Bolitoglossa bolanosi sp. nov. (A) Paratopotype male subadult UCR 22966, (B) paratype juvenile UCR 24247, (C) paratype adult female UCR 22421, and (D) paratype adult female UCR 24245. Photographs A and C by Erick Arias, B and D by Omar Ziitiga. locality of B. splendida is only ~4 km from the site with B. bolanosi. Distribution. The known distribution area of Bolitoglossa bolanosi is very small, restricted throughout to ~15 km on the summits of the peaks Durika, Arbolado, Haku, and Utyum on the Cordillera de Talamanca (Fig. 1). The altitudinal range of the new species is 2,550—3,240 m asl. All the populations of the new species were found in primary vegetation (paramo and forest) and all are within a protected area, La Amistad International Park. Conservation status. The conservation status of this species is uncertain; however, its known distribution range is small (<20,000 km?) and all know populations are restricted to summits of peaks, sites that are highly vulnerable to forest fires and other effects of climate change. We suggest that it should be tentatively considered as Vulnerable (VU) following the IUCN (2017) category criteria. Discussion Costa Rica hosts 56 species of salamanders, and it is the sixth most diverse country in the World in terms of total species richness and the most diverse relative to its total area (AmphibiaWeb 2023; Boza-Oviedo et al. 2012; Frost 2023). Several new species of salamanders have been described from ICA in this century (Arias and Kubicki Amphib. Reptile Conserv. 2018; Bolafios and Wake 2009; Boza-Oviedo et al. 2012: Garcia-Paris et al. 2008; Hanken et al. 2005; Kubicki 2016; Kubicki and Arias 2016; Kubicki et al. 2022: Wake et al. 2007). Nevertheless, the taxonomic studies of salamanders in this region have been obscured due to a conservative morphology (Arias and Kubicki 2018; Boza-Oviedo et al. 2012; Kubicki et al. 2022). Recently, Kubicki et al. (2022) found low genetic distances among species of Nototriton from Costa Rica; however, they showed that two different species of Nototriton coexist (sympatry) in two sites, providing strong evidence of speciation. We also found relatively low (Table 1) genetic distances among the species of the B. subpalmata S.G., however, we found two sites separated by less than 7 km ina straight line from where two different species occur. In the first case, in the paramo of the Cerro Durika, samples of B. bolanosi (UCR 24245-7) were found only 4 km in a straight line from specimens of B. pesrubra (UCR 23049). The area separating these two sites consisted of homogeneous paramo, lacking any significant features that could be viewed as a barrier that would prevent the potential mixture of individuals or genes. However, the genetic distance among these organisms is 2.14% for 16S, providing evidence of genetic isolation. A second case was found in the paramo of the Cerro Utyum, where the B. bolanosi specimens (UCR 22421—7) were found only 7 km ina straight line from B. kamuk (UCR 23047). Similarly, these sites are separated by homogeneous paramo and we found genetic distances of 1.35—1.8% for December 2023 | Volume 17 | Number 1 & 2 | e327 Arias et al. 16S. We acknowledge the limitations of our conclusions, since our genetic analyses were based solely on the mitochondrial results. However, we think that all the evidence supports our hypothesis about the relationships within the B. subpalmata S.G. Bolitoglossa bolanosi is the fourth species of salamander endemic to the paramo of ICA and the Montane rain forest surrounding it. The other three species also belong to the B. subpalmata S.G. Although there are relatively few species, the role of the paramos of ICA in the speciation of salamanders is interesting, especially since the paramos correspond only to 0.3% of the continental surface of Costa Rica (Kappelle and Horn 2016). The current isolated patches of paramos in ICA and its contraction-expansion dynamics in the past can explain (in part) the species formation in this habitat. In addition, it is noteworthy that the larger paramo in Costa Rica, the Cerro Chirripo, has not been sampled for salamanders. This paramo corresponds to two-thirds of all paramo in the country. There is a need for more studies 1n the paramos of ICA, especially due to the vulnerability of this ecosystem to climate change (Enquist 2002; Nogués- Bravo et al. 2007). In the past, B. pesrubra was relatively common (Bolafios and Wake 2009; Vial 1968). However, its populations have been greatly reduced (Boza-Oviedo et al. 2012; Garcia-Paris et al. 2008), and we do not fully understand the cause of this reduction. Acknowledgements.—We thank Laura Marquez- Valdelamar, Nelly Lopez, and Andrea Jiménez-Marin for their laboratory assistance; Federico Bolafios for the use of specimens from the Museo de Zoologia of the Universidad de Costa Rica; and Omar Zufiiga and Olmer Cordero who provided valuable assistance in the field during the expeditions. EA thanks the Posgrado en Ciencias Biologicas for supporting this study, the CONACYyT for a student grant (CVU/Becario) 626946/330343, and the Programa de Innovacion y Capital Humano para la Competitividad PINN-MICITT for a student grant (PED- 0339-15-2). The fieldwork was partially supported by the National Geography Society (Grant number W-346-14). The laboratory work was partially funded by Direccion General de Asuntos del Personal Académico, UNAM, under grant PAPHT-UNAM no. IN205521 to GPO. 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MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution 30: 2,725—2,729. Vial JL. 1968. The ecology of the tropical salamander, Bolitoglossa subpalmata, in Costa Rica. Revista de Biologia Tropical 15: 13-115. Wake DB. 1987. Adaptive radiation of salamanders in Middle American cloud forests. Annals of the Missouri Botanical Garden 74: 242-264. Wake DB, Savage JM, Hanken J. 2007. Montane salamanders from the Costa Rica-Panama_ border region, with descriptions of two new species of Bolitoglossa. Copeia 2007: 556-565. Zwickl DJ. 2006. Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. Ph.D. Dissertation, The University of Texas, Austin, Texas, USA. 125 p. Amphib. Reptile Conserv. 156 Erick Arias is a Costa Rican herpetologist who earned his Bachelor’s degree in Biology at Universidad de Costa Rica in 2013. Erick then joined Gabriela’s Lab at Universidad Nacional Autonoma de México (UNAM) where he received a Doctorate in Systematics in 2019 studying the systematic, taxonomy, and biogeography of the neotropical frogs in the Craugastor podiciferus species group. In 2022, Erick joined the Universidad de Costa Rica as professor and researcher. Erick’s current research focuses on the systematics, taxonomy, and biogeography of the amphibians and reptiles of Costa Rica, especially direct-developing frogs and salamanders. Gerardo Chaves is a retired biologist, associated with the Zoology Museum of the University of Costa Rica. His undergraduate thesis focused on the arrival of Olive Ridley Sea Turtles, but most of his professional work has been oriented towards the ecology and taxonomy of the Costa Rican herpetofauna. His research activity is directed towards understanding the declines of amphibian populations in Mesoamerica and filling in information gaps in the Cordillera de Talamanca. From this research, he has published articles in several journals related to the ecology and taxonomy of Neotropical herpetofauna. Gerardo is currently the Chair of the IUCN Amphibian Specialist Group in Costa Rica. Gabriela Parra Olea is a researcher at the Instituto de Biologia, UNAM, Mexico, and curator of Mexico’s national collection of amphibians and reptiles. Her research is focused on the molecular systematics and conservation of Mexican amphibians. Her laboratory is formed by students and postdocs all working on research projects in the systematics, taxonomy, conservation genetics, and impact of infectious diseases, specifically chytridiomycosis, on the conservation of amphibians. December 2023 | Volume 17 | Number 1 & 2 | e327 Arias et al. Appendix 1. Institutional voucher numbers, locality information, and GenBank accession numbers for the specimens used in the molecular phylogenetic analyses. Museum collection acronyms follow Frost (2023) with the addition of EAP to refer to Erick Arias field numbers and CRARC to refer to the Costa Rica Amphibian Research Center private collection. NV= No voucher. CR = Costa Rica, HN = Honduras, MX = Mexico. Species B. mexicana B. aurae B. sombra B. compacta B. splendida B. bramei B. bramei B. bramei B. bramei B. bramei B. bramei B. gomezi B. gomezi B. gomezi B. gomezi B. gomezi B. gomezi B. gomezi B. gomezi B. gomezi B. gomezi B. gomezi B. gomezi B. gomezi B. gracilis B. gracilis B. gracilis B. tica Amphib. Reptile Conserv. pease [owimramemer [ae fos fam | ees [aawsearmener [aw fae [ae [ oon [omsearence [ave fore [oor [ MVZ229170 Paraiso, Cartago, CR 1680 9/692 -83.782 AY526121 MVZ229171 Paraiso, Cartago, CR 1680 9.692 -83.782 AY526122 UCR20541 Paraiso, Cartago, CR 1400 9.734 -83.764 OR162558 UCR12065 Guarco, Cartago, CR 2440 9.686 -83.894 AY526137 157 December 2023 | Volume 17 | Numbe cyt b GU725470 KX779528 AY526174 JQ899193 JQ899181 AF212066 JQ899189 JQ899190 JQ899172 OR147299 OR147300 JQ899184 JQ899185 JQ899186 JQ899187 JQ899188 JQ899170 JQ899177 JQ899178 JQ899179 JQ899180 JQ899169 JQ899171 JQ899176 AF212067 AF212068 AF212089 r1&2 | e327 A new species of Bolitoglossa Appendix 1 (continued). Institutional voucher numbers, locality information, and GenBank accession numbers for the specimens used in the molecular phylogenetic analyses. Museum collection acronyms follow Frost (2023) with the addition of EAP to refer to Erick Arias field numbers and CRARC to refer to the Costa Rica Amphibian Research Center private collection. NV= No voucher. CR = Costa Rica, HN = Honduras, MX = Mexico. enne ade Collection locality Elevation (m) | Latitude | Longitude cyt b UCR20514 Paraiso, Cartago, CR 2300 9.712 -83.858 JQ899162 JQ899192 B. subpalmata | MVZ190875 Barva, Heredia, CR 1850 10.133 pwr | = | AF212093 B. subpalmata | MVZ194827 San Ramon, Alajuela, CR 1500 10.033 -84.483 AF212090 B. subpalmata | MVZ194828 San Ramon, Alajuela, CR 1500 10.033 -84.483 EU448107 AF212091 B. subpalmata | MVZ194889 Se usaed Corauade; Sah 1700 9,998 -83.964 p= | AF212095 B. subpalmata | MVZ229172 Monte Verde, Puntarenas, CR 1500 AF416697 | AF212094 iy nnaiman| SY -Bisnte Heredia, CR ND ND ND AF212092 Carrillo B. subpalmata | UCR20833 Alfaro Ruiz, Alajuela, CR 1600 10.177 | -84.391 | OR162562 Amphib. Reptile Conserv. 158 December 2023 | Volume 17 | Number 1 & 2 | e327 mi ee UCR22423 Buenos Aires, Puntarenas, CR 2870 9.323 -83.187 OR162573 | OR147309 cating UCR22422 Buenos Aires, Puntarenas, CR 2870 9.323 -83.187 | OR162571 | OR147307 Arias et al. Appendix 1 (continued). Institutional voucher numbers, locality information, and GenBank accession numbers for the specimens used in the molecular phylogenetic analyses. Museum collection acronyms follow Frost (2023) with the addition of EAP to refer to Erick Arias field numbers and CRARC to refer to the Costa Rica Amphibian Research Center private collection. NV= No voucher. CR = Costa Rica, HN = Honduras, MX = Mexico. Species eran att Collection locality Elevation (m) | Latitude | Longitude cyt b ss eee UCR24245 Buenos Aires, Puntarenas, CR 3240 9.374 -83.303 | OR162568 | OR147304 a on = =a AF212075 Amphib. Reptile Conserv. 159 December 2023 | Volume 17 | Number 1 & 2 | e327 A new species of Bolitoglossa Appendix 2. Specimens examined in the morphological analysis. All voucher numbers below are ‘UCR’ numbers, and the specimens were housed at the Museo de Zoologia at the Universidad de Costa Rica. Bolitoglossa bolanosi sp. nov. COSTA RICA: Limon, Talamanca, Cerro Utyum: (224212); Puntarenas, Buenos Aires, Cerro Arbolado: (22965); Puntarenas, Buenos Aires, Cerro Haku: (242459, 24246); Puntarenas, Buenos Aires, Cerro Haku: (22741-4°, 227453); Puntarenas, Buenos Aires, Cerro Utyum: (224224, 224239, 22424, 224259). Bolitoglossa kamuk COSTA RICA: Limon, Talamanca, Cerro Dika: (23047); Limon, Talamanca, Cerro Kamuk: (208523). Bolitoglossa pesrubra COSTA RICA: Cartago, El Guarco, Cerro de la Muerte: (23912, 224369); Cartago, Paraiso, Cerro de la Muerte: (138106, 19034, 1912592, 190999, 23634); San José, Dota, Cerro de la Muerte: (120674, 207932, 20797-92); San José. Pérez Zeled6on, Cuerici: (117124, 117574, 12109’). Amphib. Reptile Conserv. 160 December 2023 | Volume 17 | Number 1 & 2 | e327 Official journal website: amphibian-reptile-conservation.org Lan Amphibian & Reptile Conservation A 17(1 & 2) [Taxonomic Section]: 161-173 (e328). urn:lsid:zoobank.org:pub:FB17AF57-AAE5-4668-8D22-1722DAEF4AA5 Two new species of gymnophthalmid lizards of the genus Petracola (Squamata: Cercosaurinae) from the Andes of northeastern Peru, and their phylogenetic relationships 12,3."Luis Mamani, *°Victor J. Vargas, ?*°Juan C. Chaparro, and °Alessandro Catenazzi 'Programa de Doctorado en Sistemdatica y Biodiversidad, Facultad de Ciencias Naturales y Oceanogrdaficas, Universidad de Concepcion (UdeC), Concepcion, CHILE *Museo de Historia Natural de la Universidad Nacional de San Antonio Abad del Cusco (MHNC), Plaza de Armas s/n (Paraninfo Universitario), Cusco, PERU *Museo de Biodiversidad del Pern (MUBI), Cusco, PERU *Asociacion Pro Fauna Silvestre Ayacucho, Huamanga, Ayacucho, PERU °Servicio Nacional Forestal y de Fauna Silvestre (SERFOR), Lima, PERU ‘Department of Biological Sciences, Florida International University (FIU), Miami, Florida 33199, USA Abstract.—The small and semi-fossorial lizards of the genus Petracola are distributed in the Andes from central to northern Peru, on both sides of the dry valley of the Maranon River. Very little is known about the taxonomy of Petracola, but it is presumed that this genus harbors a higher number of species than is currently known. Here, we used a Maximum Likelihood (ML) molecular phylogenetic analysis using concatenated fragments of four mitochondrial genes (12S, 16S, cyt-b, and ND4) and one nuclear gene (c-mos), and the revision of museum specimens, to determine the species diversity of populations of Petracola on both sides of the Maranon River. Our results show that Petracola is a monophyletic genus composed of three independent lineages. The first lineage includes two undescribed species from the right side of the Maranon River, which we describe herein. The second lineage corresponds to P. ventrimaculatus, and the third lineage includes P. waka which contains undescribed species. Among the two new species from the right side of the Maranon River, P. amazonensis can be identified by having only the first superciliary expanded to the dorsal surface, two pairs of genials in contact, absence of a loreal scale, venter orange with black spots forming a transverse band, absence of precloacal pores, and a maximum SVL of 43.0 mm in females; whereas P. shurugojalcapi can be identified by having two genials in contact, absence of a loreal scale, two discontinuous superciliaries, black venter, dorsum dark brown or black with some cream spots, and an absence of precloacal pores. The two new species increase the diversity of this genus to seven species, four of which inhabit the right side of the dry valley of the Maranon River. However, like other genera of gymnophthalmid Andean lizards, the diversity of Petracola is still underestimated. Keywords. Andean lizard, Cercosaurini, dry valley, endemic species, northern Andes, Marafion River, sem1-fossoriality Citation: Mamani L, Vargas VJ, Chaparro JC, Catenazzi A. 2023. Two new species of gymnophthalmid lizards of the genus Petracola (Squamata: Cercosaurinae) from the Andes of northeastern Peru, and their phylogenetic relationships. Amphibian & Reptile Conservation 17(1 & 2) [Taxonomic Section]: 161-173 (e328). Copyright: © 2023 Mamani et al. This is an open access article distributed under the terms of the Creative Commons Attribution License [Attribution 4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. The offcial and authorized publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org. Accepted: 4 October 2023; Published: 30 December 2023 Introduction The systematics and taxonomy of Petracola are poorly developed, and geographic information gaps prevent an The genus Petracola Doan and Castoe, 2005, isa poorly —_ appreciation of the full diversity and distribution of the studied Andean lineage of small and semi-fossorial lizards distributed in the central and northern Andes of Peru, on both margins of the dry valley of the Marafion River (DVMR) (Doan and Castoe 2005; Kizirian et al. 2008; Kohler and Lehr 2004; Uzzell 1970). The species of Petracola inhabit ecosystems of montane cloud forests, inter-Andean valleys, and wet puna at elevations from 1,889 to 3,600 m asl. Correspondence. luismamanic@gmail.com Amphib. Reptile Conserv. genus. The type species of Petracola, P. ventrimaculatus (Boulenger, 1900), was initially assigned to Proctoporus by Boulenger (1900). Previously, based on morphological characters such as the presence of smooth dorsal scales, three postparietal scales, short limbs, and the presence of entire palpebral disc, Uzzell (1970) assigned P. ventrimaculatus to the Proctoporus pachyurus group. Further analyses based on molecular data and a taxonomic December 2023 | Volume 17 | Number 1 | e328 Two new species of Petracola from Peru review led to the creation of the genus Petracola (Castoe et al. 2004; Doan and Castoe 2005), which has been supported by recent studies (Goicoechea et al. 2012; Mamani et al. 2020, 2022; Moravec et al. 2018; Torres-Carvajal et al. 2016). Species of Petracola are phylogenetically related to Andean speciose linages containing Cercosaura, Potamites, and Proctoporus, and minor lineages such as “Cercosaura manicata_ boliviana,’’ Dendrosauridion, Selvasaura, and Wilsonosaura (Mamani et al. 2020, 2022; Moravec et al. 2018; Rojas-Runjaic et al. 2021). Despite the increasing knowledge about high-Andean gymnophthalmids accumulated over the past decade, the diversity of gymnophthalmid species continues to be underestimated. Currently, the genus contains five species: Petracola angustisoma Echevarria and Venegas, 2015; P labioocularis (Kohler and Lehr, 2004); P pajatensis Rodriguez and Mamani, 2020; P. ventrimaculatus, and P waka, Kizirian, Bayefsky-Anand, Eriksson, Le and Donnelly 2008. Although Kizirian et al. (2008) hypothesized that P. ventrimaculatus and P. waka represent complexes of species, only two more species have been described since 2015 (Echevarria and Venegas 2015; Rodriguez and Mamani 2020). Moreover, this genus has a peculiar distribution on both sides of the DVMR, in which Petracola angustisoma and P. pajatensis are distributed on the right side (Echevarria and Venegas 2015; Kohler and Lehr 2004; Rodriguez and Mamani 2020), P waka is distributed on the left side, and P. ventrimaculatus 1s distributed on both sides of the Marafion River (Kizirian et al. 2008). Vicariant topographic features such as the DVMR introduce genetic barriers between populations on the two sides of the valley, promoting the diversification of lineages (Futuyma and Kirkpatrick 2017). The ornithological literature supports the hypothesis that the arid valley of the DVMR interrupts gene flow between populations and promotes allopatric speciation (Hazzi et al. 2018; Weir 2009; Winger and Bates 2015), but this hypothesis has not been tested in other biological groups, and especially in philopatric organisms such as gymnophthalmid lizards (Doan et al. 2021). Here, based on an analysis of morphological data and molecular phylogenetic relationships of four mitochondrial genes (12S, 16S, cyt-b, and ND4) and one nuclear gene (c-mos) from specimens collected on both sides of the DVMR, we evaluated the hypothesis proposed by Kizirian et al. (2008) that Petracola ventrimaculatus and P. waka are species complexes, and we describe two new species from those complexes. Material and Methods Data Collection and Morphology Specimens of the new species were collected in Department Amazonas, Peru, and are deposited in the Coleccién Cientifica Pro Fauna Ayacucho (PFAUNA) and the Museo de Biodiversidad del Pert (MUBI). Specimens were collected by hand, euthanized, fixed in 10% formalin, and Amphib. Reptile Conserv. stored in 70% ethanol. Tissue samples were stored in 2 ml cryogenic tubes filled with 96% ethanol. The terminology for diagnostics and format description follows Uzzell (1970), Kizirian (1996), Mamani et al. (2022), and Mamani and Rodriguez (2022). Measurements were taken with calipers to the nearest 0.1 mm. Abbreviations for measurements are as follows: SVL (snout-vent length), TL (tail length), LAL (length between arm and leg), LSA (length from the tip of snout to anterior margin of the insertion of the arm), HL (head length, from the tip of snout to posterior margin of tympanum), HW (head width, maximum width of head), HH (head height, maximum height of head), FR (frontal length), and FN (frontonasal length). Morphological data of Petracola species were taken from the literature (Echevarria and Venegas 2015; Kizirian et al. 2008; Kohler and Lehr 2004; Rodriguez and Mamani 2020) and the examination of voucher specimens deposited at the Museo de Historia Natural de la Universidad Nacional de San Marcos (MUSM), Lima, Peru; Museo de Biodiversidad del Peru (MUBI), Cusco, Peru; and Coleccion Cientifica Pro Fauna Silvestre (PFAUNA), Ayacucho, Peru. The specimens examined are listed in Appendix I. DNA Extraction, Amplification, Sequencing, and Phylogenetic Reconstruction We extracted DNA from the tissue samples of seven voucher specimens of Petracola (Table 1) from both sides of the DVMR. We used standard protocols (Mamani et al. 2020) to amplify fragments of the small subunit rRNA (12S), large subunit rRNA (16S), dehydrogenase subunit 4 (ND4), cytochrome b (cyt-b), and oocyte maturation factor gene (c-mos). The new DNA sequences are available in GenBank (Appendix 1; https://www.ncbi.nlm. nih.gov/genbank/). We obtained additional sequences of congeneric species and representative species of different gymnophthalmid genera and outgroups (following Moravec et al. 2018) from GenBank. We excluded Petracola ventrimaculatus (AY 507894) from our data set of the gene ND4, because the sample was nested within Cercosaura genus in a previous exploratory analysis, suggesting a possible labeling error or contamination. The final data set contained 137 operational taxonomic units (OTU) and 2,384 bp of concatenated fragments (458 bp for 12S, 561 bp for 16S, 345 bp for cyt-b, 639 bp for ND4, and 381 bp for c-mos) (Table S1). The genetic sequences were aligned using MUSCLE software (Edgar 2004) implemented in MEGA-X (Kumar et al. 2018). We inferred the phylogenetic relationships using a Maximum Likelihood (ML) approach in IQTREE Web server (Trifinopoulos et al. 2016). The evolution models were estimated in ModelFinder (Kalyaanamoorthy et al. 2017) and were TIM2+F+1+Gé4 for 12S, GTR+F+R4 for 16S, TIM2e+I+Gé4 for cytb, GIR+F+R5 for ND4, and TPM3+F+G4 for c-mos. Branch supports were estimated for 10,000 replicates using ultrafast Bootstrap (Hoang et al. 2018). Following Moravec et al. (2018), we used Alopoglossus atriventris, Bachia bresslaui, B. dorbignyi, B. flavescens, Ecpleopus gaudichaudii, Gymnophthalmus December 2023 | Volume 17 | Number 1 | e328 Mamani et al. Table 1. Voucher museum specimens of Petracola lizards, mentioning their collection locations and GenBank codes sequences used in this study. All localities are from Peru. Species/voucher Locality Chiliquin, Chachapoyas, P. amazonensis MUBI 11473 Amazonas La Jalca, Chachapoyas, P. shurugojalcapi MUBI 17727 Amazonas La Jalca, Chachapoyas, P. shurugojalcapi PFAUNA 430 Amazonas Bafios del Inca, P. waka MUBI 2603 , Cajamarca, Cajamarca Bafios del Inca, P. waka MUBI 2605 ; Cajamarca, Cajamarca Type locality, Cajabamba, P. waka MUBI 2609 ; Cajamarca Type locality, Cajabamba, P. waka MUBI 2611 Cajamarca leucomystax, Rhachisaurus brachylepis, | Riolama inopinata, and R. leucosticta as outgroup taxa. We estimated uncorrected p-distances between species of Petracola for the 16S gene using MEGA-X (Kumar et al. 2018). Genetic distances for 16S larger than 3% separate most species of gymnophthalmid lizards (Torres- Carvajal et al. 2015). The electronic version of this article in Portable Document Format (PDF) will represent a published work according to the International Commission on Zoological Nomenclature (ICZ), and hence the new name contained in the electronic version is effectively published under the Code from the electronic edition alone. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved, and the associated information viewed through any standard web browser by appending the LSID to the prefix http://zoobank.org/. The LSID for this publication is: urn:lsid:zoobank.org:pub:FB17AF57- AAE5-4668-8D22-1722DAEF4AAS. Results Phylogenetic and Generic Assignment Relationships The ML tree recovered the monophyly of the ingroup Cercosaurinae (ultrafast bootstrap, UB = 100; Figs. 1 and S1). Our topology recovered the monophyly of all genera with high support, except Proctoporus (UB = 52). The monophyly of Oreosaurus was not supported, because O. serranus was recovered as a different lineage of Oreosaurus, with Cercosaura, “Cercosaura manicata boliviana,’’ Dendrosauridion, Petracola, Potamites, Proctoporus, and Wilsonosaura as its closest relatives. The genus Petracola was recovered with high support (UB = 100) as sister lineage of Cercosaura, “Cercosaura Amphib. Reptile Conserv. OR231541 OR231542 OR231543 OR231544 OR231545 OR231546 OR231547 12S 16S ND4 cytb c-mos OR231652 OR208583 OR198057 OR211561 OR231653 - OR198058 OR211562 OR231654 - OR198059 = OR211563 OR231655 OR208584 OR198060 OR211564 OR231656 OR208585 OR198061 OR211565 OR231657 OR208586 OR198062 - OR231658 OR208587 OR198063 OR211566 manicata boliviana,’”’ Dendrosauridion, Potamites, Proctoporus, Selvasaura, and Wilsonosaura (Fig. 1). The specimens of Petracola from the left side of the DVMR were monophyletic, and include P. ventrimaculatus and P. waka, but the node support was low (UB = 83). Three specimens from the right side of the DVMR, including the two new species (P. amazonensis and P. shurugojalcapi), were recovered as sister lineages with high support (UB = 100) and have high uncorrected p-distances for the 16S gene ranging from 3.7% to 8.0% (Table 2), suggesting they might be distinct from the currently recognized taxa. Additionally, the distance between specimens of P. waka from the type locality (MUBI 2609, 2611) and those from Bafios del Inca and Abra Gelic is 6.3-6.6%, which suggests a hidden cryptic diversity (Table 2). Finally, our phylogeny and morphological traits such as presence of imbricate and scale-like papillae on the tongue, scales of the head without striations or rugosities, smooth dorsal scales, and absence of prefrontal scales (Doan and Castoe 2005), support the assignment to the genus Petracola. Taxonomy Petracola amazonensis sp. nov. urn:Isid:zoobank.org:act:89A5D519-171D-4906-B82B-6C60CB8F9854 Holotype. MUBI 11485, adult female (Figs. 2-4) from Upa, bridle path to Yurumarca, District of Chiliquin, Province of Chachapoyas, Department of Amazonas, Peru (6°0’9.19” S; 77°49’21.08” W; ca. 3,020 m asl), collected by Juan C. Chaparro on 23 May 2012. Paratypes. Two specimens: A subadult male (MUBI 11473) and a subadult female (MUBI 11474) from near the type locality (5°59’40.27” S; 77°48’36.26” W; 3,250 m asl), collected by Juan C. Chaparro and Alexander Pari on 22 May 2012. December 2023 | Volume 17 | Number 1 | e328 Two new species of Petracola from Peru 100 Neusticurus Placosoma Kataphraktosaurus ‘| Echinosaura Centrosaura Riama Gelanesaurus Pholidobolus { Rheosaurus Oreosaurus Oreosaurus serranus Cercosaura Potamites Cercosaura manicata boliviana Selvasaura Dendrosauridion Proctoporus Wilsonosaura Petracola waka MUBI 2611 Petracola waka MUBI 2609 Petracola waka MUBI 2605 Petracola waka MUBI 2603 Petracola waka KU212687 Petracola ventrimaculatus KU219838 Petracola ventrimaculatus CORBIDI 9235 Petracola amazonensis MUBI 11473 Petracola shurugojalcapi MUBI 17727 Petracola shurugojalcapi PFAUNA 430 {00 Fig. 1. Phylogenetic relationships of Cercosaurini (log likelihood = -54268.289, ultrafast bootstrap = 10,000) constructed from the data set of 2,384 nucleotides for mitochondrial genes (12S, of Petracola and the two new species Petracola amazonensis bootstraps values. 16S, cyt-b, and ND4) and a nuclear gene (c-mos), showing species and P. shurugojalcapi. The numbers on the branches are ultrafast Table 2. Pairwise uncorrected p-distances for 16S rRNA between Petracola species. The asterisk (*) indicates type locality. 1 2 (1) P. ventrimaculatus CORBIDI 9235 - (2) P. ventrimaculatus KU 219838 0.024 - (3) P. waka KU 212687 0.063 0.071 (4) P. waka MUBI 2603 0.073 0.091 (5) P. waka MUBI 2605 0.073 0.091 (6) P. waka MUBI 2609* 0.069 0.082 (7) P. waka MUBI 2611* 0.069 0.082 (8) P. shurugojalcapi MUBI 17727 0.058 0.074 (9) P. shurugojalcapi PFAUNA 430 0.058 0.074 (10) P. amazonensis MUBI 11473 0.057 0.072 Etymology. The specific epithet, amazonensis, 1s an adjective in reference to the type locality in the Department of Amazonas, northern Peru. Diagnosis: Petracola amazonensis is diagnosed based on the following combination of characters: (1) frontonasal longer than frontal; (2) nasoloreal suture absent; (3) two supraoculars; (4) one superciliary scale expanded on dorsal surface of head; (5) two postoculars; (6) palpebral disc transparent, entire or divided vertically in two; (7) four supralabials anterior to the posteroventral angle of the subocular; (8) 3—4 anterior infralabials; (9) four Amphib. Reptile Conserv. 164 3 4 ) 6 it 8 9 10 0.063 - 0.063 0.000 - 0.066 0.031 0.031 - 0.066 0.031 0.031 0.000 - 0.080 0.079 0.079 0.079 0.079 > 0.080 0.079 0.079 0.079 0.079 0.000 - 0.085 0.078 0.078 0.072 0.072 0.037 0.037 - genials in contact; (10) three rows of pregulars; (11) dorsal body scales rectangular, smooth, juxtaposed; (12) 25-28 scales around midbody; (13) 31-32 transverse dorsal rows; (14) 18-19 transverse ventral rows; (15) 17-20 longitudinal dorsal rows; (16) 8—9 longitudinal ventral rows; (17) a continuous series of small lateral scales separate dorsals from ventral scales; (18) 2-4 posterior cloacal plate scales; (19) two anterior preanal plate scales; (20) four femoral pores per hind limb in males, two in females; (21) preanal pores absent; (22) 7-9 subdigital lamellae on finger IV; 11-14 subdigital lamellae on toe IV; (23) limbs not overlapping when December 2023 | Volume 17 | Number 1 | e328 Mamani et al. adpressed against body; (24) pentadactyl, digits clawed; and (25) coloration of female in life: dorsum is brown with numerous dark spots distributed irregularly from tip of head to tail, flanks are similar to dorsum, the chin and throat are pale orange with large black spots, venter mostly orange, with black spots on anterior part of ventral scales forming longitudinal lines (Fig. 4); coloration of males in life is unknown; in preservative the dorsum is similar to coloration in life, and venter changes from orange to cream (Fig. 2). Petracola amazonensis can be distinguished from P. angustisoma and P. pajatensis by lacking a loreal scale (loreal scale present in P angustisoma and P. pajatensis);, from P. labioocularis by lacking precloacal pores and posterior subocular is not elongated downward (precloacal pores present and posterior subocular is elongated downward and separates supralabials in P. labioocularis), from P. waka by having two genials in contact, first superciliary scale, and venter is orange with black spots forming transverse bands (three genials in contact, four continuous superciliary scales, and venter is cream with some small, black spots in P. waka); from P. ventrimaculatus by having a maximum SVL in males of 43.0 mm, dorsum dark brown with some black spots not forming bands, and first superciliary only (maximum SVL in females 59.0 mm, dorsum light brown with continuous black dorsal bands, and 2-3 discontinuous superciliaries in P. ventrimaculatus). Description of the holotype. Adult female, snout— vent length (SVL) 43.0 mm, tail length 42.3 mm (regenerated); head scales smooth, without striations or rugosities; rostral scale wider (1.5 mm) than tall (0.8 mm), in contact with frontonasal, nasals, and first supralabials; frontonasal longer (2.5 mm) than wide (1.7 mm), longer than frontal scale, widest in the middle, in contact with rostral, nasal, first superciliary, and frontal; prefrontal absent; frontal longer that wide, pentagonal, Fig. 2. Holotype of Petracola amazonensis, female MUBI 11485 (SVL = 43.0 mm, TL = 42.3 mm). Amphib. Reptile Conserv. in contact with first supraocular and frontoparietals; frontoparietal paired, polygonal (hexagonal), in contact with frontal, supraoculars, parietals, and interparietals;: two supraoculars, in contact with superciliaries, frontal, frontoparietals, interparietal, and postoculars; parietals polygonal (irregular heptagon), in contact with frontoparietals, posterior supraocular, postocular, interparietal, temporals, supratemporals laterally, and with postparietals posteriorly; interparietals hexagonal, in contact with frontoparietals anteriorly, parietals laterally, and with postparietal posteriorly; two polygonal postparietals, joined in the middle by a small surface. Nasal scale entire, longer than high, in contact with first supralabial; loreal scale absent; left side with first superciliary, on right side with a posterior rudimentary posterior superciliary, first superciliaries expand on dorsal surface of head; two small preoculars; frenocular trapezoidal in contact with second supralabial; palpebral disc entire and transparent; three suboculars on right side and two on left side; two postoculars; temporals and supratemporals smooth, polygonal; four supralabials anterior to the posteroventral angle of the suboculars. Mental wider than long, in contact with the first infralabial and postmental posteriorly; postmental single, polygonal (irregular pentagonal), in contact with first and second infralabials, and first pair of genials; four genials in contact; three transversal rows of pregular scales, in the anterior row they are large; five rows of gular scales, quadrangular and circular. Dorsal scales rectangular with blunt edges, juxtaposed, smooth, 32 transverse rows; 17 longitudinal dorsal rows at midbody; a continuous series ip = —— Fig. 3. Drawings of lateral, dorsal, and ventral views of the head of the holotype (MUBI 11485) of Petracola amazonensis. December 2023 | Volume 17 | Number 1 | e328 Two new species of Petracola from Peru of small lateral scales; reduced scales at limb insertion region; 18 transverse ventral rows; eight longitudinal ventral rows at midbody; anterior and posterior preanal plate paired; scales on the tail rectangular, subimbricate, and smooth; ventral scales quadrangular, juxtaposed, and smooth. Limbs pentadactyl; digits clawed; dorsal brachial scales polygonal, imbricate, and smooth with blunt edges; ventral brachial scales small, rounded, and smooth; dorsal antebrachial scales polygonal, imbricate and smooth; ventral antebrachial scales rounded, smooth, smaller than dorsal; dorsal manus scales polygonal, smooth, imbricate; palmar scales small, rounded, and domelike; dorsal scales on fingers smooth, quadrangular, imbricate, two on finger I, three on II, five on III, five on IV, and four on V; two subdigital lamellae on finger I, five on finger IT, seven on finger III, seven on finger IV, five on finger V; scales on anterodorsal surface of thigh polygonal, smooth, imbricate; scales on posterior surface of thigh small, rounded, and separated; scales on ventral surface of thighs polygonal with blunt edges, smooth, imbricate, polygonal and juxtaposed; two femoral pores on each thigh; scales on anterior surface of crus polygonal, smooth, imbricate, decreasing in size distally; scales on posterodorsal surface of crus smooth, polygonal, and imbricate; scales on ventral surface of crus polygonal, enlarged, smooth, and imbricate; scales on dorsal surface of feet polygonal, smooth, and imbricate; scales on ventral surface of feet rounded, small, and domelike; dorsal scale of toes smooth, imbricate, two on toe I, four on toe II, five on toe III, six on IV, five on toe V; four subdigital lamellae on toe I, seven on toe II, nine on toe III, 13 on toe IV, eight on toe V. Coloration in life was described in diagnosis; in preservative, dorsum, dorsal surface of head, neck, arms, and legs brown with irregular, black spots; venter, chin, neck, arms, and legs gray with black spots, on venter black spots form transverse bands (Fig. 4). ae fn es * a . , : os ee Fig. 4. Holotype in life of Petracola amazonensis from Upa (MUBI 11485, SVL= 43.0 mm, TL = 42.3 mm, female). Amphib. Reptile Conserv. Variation. The most notable variation was the number of infraoculars: adult female with 2-3 (left-right), juvenile male with 1—2, and juvenile female with 3—3; adult female with a rudimentary posterior superciliar on the right side; juvenile male with three anterior infralabials on the right side and three postparietals; and adult female with 25 scales around body and 28 on juveniles. Distribution and ecology. Petracola amazonensis is only known from the type locality, Upa near Yurumarca, from 3,020-3,250 m asl (Fig. 5), District of Chiliquin, Province of Chachapoyas, Department of Amazonas (Fig. 6). All specimens were found in grassland during March (rainy season). The specimen MUBI 11473 was found under rocks, MUBI 11474 inside bromeliads of the genus 7illandsia, and MUBI 11485 walking on grassland near a creek and a forest. Petracola amazonensis is likely diurnal and semifossorial. Syntopic Squamata species include Stenocercus orientalis. Tree vegetation 1s dominated by Podocarpus (Podocarpaceae), A/nus (Betulaceae), Weinmannia (Cunnoniaceae), Ceroxylon (Arecaceae) and Hyeronima (Euphorbiaceae). Dominant species in the shrub layer include Macrocarpaea (Gentianaceae), Munnozia (Asteraceae), Chusquea (Poaceae), and Piper (Piperaceae); and herbs Anthurium (Araceae), Tillandsia (Bromeliaceae), orchids (Orchidaceae) and Peperomia (Piperaceae). Conservation. Petracola amazonensis was found in a small area in the district of Chiliquin (Fig. 5), and we suspect it might be endemic to the Department of Amazonas. The type locality is 2.5—3.0 km from the Private Conservation Area “Comunal San Pablo - Catarata Gocta,” and it is very likely that the species 1s distributed within this conservation area. In general, the habitat where we conducted our herpetological surveys was in an acceptable state of conservation in May 2012. Given the lack of knowledge regarding its geographic distribution, we recommend that P. amazonensis be categorized as Data Deficient in the Red List of the International Union for Conservation of Nature (IUCN 2022). hi Mae at = Y “a. Fig. 5. Type locality of Petracola amazonensis, Upa, District of Chiliquin, Province of Chachapoyas, Department of Amazonas. December 2023 | Volume 17 | Number 1 | e328 Mamani et al. 81°0'O"W 78°0'O"W 75°0'O"W 72°0'0"W 0°0'0" q 0°0'0" N cand 3°0'0"S 6°0'0"S Brazil 9°0'0"S Pacific Ocean 0 62.5125 250 12°0'0"S 12°0'0". 81°0'O"W 78°0'O"W 75°0'O"W 72°0'0"W Fig. 6. Map showing the type localities of Petracola species: Fig. 7. Holotype of Petracola shurugojalcapi, adult female Petracola amazonensis (blue cross), P. angustisoma (red PFAUNA 431 (SVL = 51.0 mm, TL = 39.0 mm). circle), P. labioocularis (light blue triangle), P. pajatensis (green pentagon), P. shurugojalcapi (yellow star), P. ventrimaculatus (purple cross), and P. waka (orange diamond). (2D iG ey, ae ae | Cl © eal eas aug \\ ee l} ai eee — Fig. 9. Holotype in life of Petracola shurugojalcapi from La Jalca Grande (PFAUNA 431, SVL = 51.0 mm, TL = 39.0 mm, adult female). Fig. 8. Drawings of lateral, dorsal, and ventral views of the head of the holotype (PFAUNA 431) of Petracola shurugojalcapi. Amphib. Reptile Conserv. 167 December 2023 | Volume 17 | Number 1 | e328 Two new species of Petracola from Peru Petracola shurugojalcapi sp. nov. urn:Isid:zoobank.org:act:2C3D891D-C75F-4B48-8951-F7870CCE4A09 Holotype. PFAUNA 431, adult female (Figs. 7—9) from Area de Conservacion Privada Llamapampa-La Jalca, District of Jalca Grande, Province of Chachapoyas, Department of Amazonas, Peru (6°25’36” S:; 77°45’56” W; 2,990 m asl), collected by Victor Vargas on 30 May 2013. Paratypes. Seven specimens: Two adult males (PFAUNA 427, MUBI 17727), subadult male (PFAUNA 432), and two adult female (PFAUNA 429, MUBI 17726) from the same place as the holotype. A male (PFAUNA 430) and female (PFAUNA 433) were collected near the type locality (6°25’36” S; 77°45’56” W; ca. 2,990 m asl). Etymology. The specific epithet, shurugojalcapi, treated as a noun in apposition, is a combination of two local words: “shurugo” the local name for a gymnophthalmid lizard and “jalcapi” that means “from Jalca” in the Quechua language. The specific name was proposed by the inhabitants of La Jalca Grande. Diagnosis. Petracola shurugojalcapi is diagnosed based on the following combination of characters: (1) frontonasal longer than frontal; (2) nasoloreal suture absent; (3) two supraoculars; (4) two discontinuous superciliaries, first expanded onto dorsal surface of head; (5) two postoculars; (6) palpebral disc transparent, divided vertically in two; (7) 3-4 supralabials anterior to the posteroventral angle of the subocular; (8) four anterior infralabials; (9) four genials in contact; (10) three rows of pregulars; (11) dorsal body scales rectangular, smooth, juxtaposed; (12) 29-32 scales around midbody; (13) 31-33 transverse dorsal rows; (14) 18—21 transverse ventral rows; (15) 19-23 longitudinal dorsal rows; (16) eight longitudinal ventral rows; (17) a continuous series of small lateral scales separating dorsals from ventrals; (18) 2-4 posterior cloacal plate scales; (19) two anterior preanal plate scales; (20) 5—6 femoral pores per hind limb in males, 2—3 in females; (21) preanal pores absent; (22) 8—9 subdigital lamellae on finger IV; 13-15 subdigital lamellae on toe IV; (23) limbs not overlapping when adpressed against body; (24) pentadactyl, digits clawed; (25) coloration of males in life is brown or dark brown with or without numerous cream spots distributed irregularly on flanks, and dorsum forming four discontinuous transversal lines, venter immaculate black or blackish gray; females with brown dorsum, numerous and irregular light brown or cream spot on flanks, venter blackish gray. Petracola shurugojalcapi can be distinguished from P. amazonensis by having dorsum brown with irregular dark spots, venter black with lateral cream spots, and two separate superciliaries (dorsum brown or dark-brown with irregular cream spots, venter orange with black spots forming transversal bands, and only first superciliary present in P. amazonensis), from P. angustisoma and P. pajatensis by not having a loreal scale (loreal scale present Amphib. Reptile Conserv. in P. angustisoma and P. pajatensis), from P. labioocularis by lacking precloacal pores and by having the posterior subocular not elongated downward (precloacal pores present and by having posterior subocular scale elongated downward and separates supralabials in P. lJabioocularis), from P. waka by having the palpebral disc divided vertically, two genial scales in contact, two discontinuous superciliary scales, venter black (palpebral disc entire, three genials in contact, four continuous superciliary scales, venter cream with small black spots in P. waka); from P. ventrimaculatus by having a maximum SVL in males of 51.0 mm, dorsum dark brown or black with some cream spots not forming bands, and venter dark with lateral cream spots (maximum SVL in males 71.1 mm, dorsum light brown with continuous black longitudinal bands, and venter cream with bold black transversal band in P. ventrimaculatus). Description of the holotype. Adult female, snout—vent length (SVL) 51.0 mm, tail length 39.0 mm (regenerated), head scales smooth, without striations or rugosities; rostral scale wider (2.0 mm) than tall (0.9 mm), in contact with frontonasal, nasals, and first supralabials; frontonasal longer (2.5 mm) than wide (2.1 mm), longer than frontal scale, widest in the middle part, in contact with rostral, nasal, first superciliary, and frontal; prefrontal absent; frontal longer than wide, pentagonal, in contact with first superciliary, first supraocular, and frontoparietals; frontoparietal paired, polygonal (hexagonal), in contact with frontal, supraoculars, parietals, and interparietals; supraoculars two, in contact with superciliaries, frontal, frontoparietals, interparietal, and postoculars; parietals longer than wide, polygonal (irregular heptagon), in contact with frontoparietals and superior supraocular, interparietal, temporals, and supratemporals laterally, and with postparietals posteriorly; interparietals polygonal (irregularly heptagonal), in contact with frontoparietals anteriorly, with parietals laterally, and with postparietal posteriorly; three postparietals, the middle one is smaller than laterals, polygonal. Nasal scale entire, longer than high, in contact with first and second supralabials; nasal suture does not divide the nasal, the right side has upper and lower suture, but the left side only the lower; loreal scale absent; two superciliaries, discontinuous, and first expanded onto dorsal surface of head; two preoculars; frenocular trapezoidal in contact with second and third supralabial; palpebral disc transparent and divided in two; three suboculars; two postoculars; temporals smooth, polygonal; four supralabials anterior to the posteroventral angle of suboculars. Mental wider than long, in contact with first infralabials and postmental posteriorly: postmental single, polygonal (irregular heptagonal), in contact with the first and second infralabials, and the first pair of genials; four genials in contact; three transversal rows of pregular scales; six gular scale rows, quadrangular with rounded corners, and smooth. Dorsal scales rectangular, longer that wide, juxtaposed, smooth, 34 transverse rows; 20 longitudinal dorsal scale rows at midbody; a continuous series of small lateral scales; reduced scales at limb insertion region present; 20 December 2023 | Volume 17 | Number 1 | e328 Mamani et al. Table 3. Morphometric measurements of Petracola amazonensis and P. shurugojalcapi. * broken tail, ** regenerated tail. P. shurugojalcapi P. amazonensis PFAUNA 431 PFAUNA 427 PFAUNA 430 PFAUNA 429 MUBI 17727 MUBI 11485 Holotype Paratype Paratype Paratype Paratype Holotype Sex Female Male Male Female Male Female SVL 51.0 48.5 440 A474 50.8 43.0 LAL 26.7 24.5 24.6 26.3 26.2 21.6 LSA 155 19 15 14.8 18.2 14.8 TL 39.0 (**) 65.9 (**) 529 PALF) 324A(F%) 42.3 HL 9.3 10.7 8.9 8.4 10.5 9.3 HW 6.8 8.2 6.9 6.2 8.1 5.6 HH S23 6.3 5.4 5.1 6.1 46 FR 1.9 2.0 1.9 1.4 2 1.4 FN 2.6 De] 2.1 Di 2.6 Di transverse ventral scale rows; eight longitudinal ventral scale rows at midbody, lateral scales slightly smaller; anterior and posterior preanal plate scales paired; scales on tail rectangular, slightly subimbricate, and smooth; ventral scales quadrangular, juxtaposed, and smooth. Limbs pentadactyl; digits clawed; dorsal brachial scales polygonal, imbricate, and smooth; ventral brachial scales rounded, separate, and smooth; dorsal antebrachial scales polygonal, imbricate and smooth; ventral antebrachial scales rounded, separate, smooth, smaller than dorsal; dorsal manus scales polygonal, smooth, imbricate; palmar scales small, rounded, separate, and dome-like; dorsal scales on fingers smooth, quadrangular, imbricate, two on finger I, five on II, five on III, five on IV, and four on V; five subdigital lamellae on finger I, eight on finger II, nine on finger IT, nine on finger IV, seven on finger V; scales on anterodorsal surface of thigh polygonal, smooth, imbricate; scales on posterior surface of thigh small, rounded, and separated; scales on anteroventral surface Fig. 10. Type locality of Petracola shurugojalcapi, Area de Conservacion Privada Llamapampa-La Jalca, District of la Jalca Grande, Province of Chachapoyas, Department of Amazonas. Amphib. Reptile Conserv. of thigh polygonal, smooth, imbricate and posteroventral surface of thigh small, polygonal and juxtaposed; two femoral pores in both thighs; scales on anterior surface of crus polygonal, smooth, separated, decreasing in size distally; scales on posterodorsal surface of crus smooth, polygonal, separated; scales on ventral surface of crus polygonal, enlarged, smooth, and imbricate; scales on dorsal surface of toes polygonal, smooth, and imbricate; scales on ventral surface of toes rounded, small and domelike; dorsal scale of toes smooth, imbricate, two on toe I, five on toe I, six on toe III, eight on IV, seven on V; five subdigital lamellae on toe I, eight on toe I, 11 on toe III, 14 on toe IV, ten on toe V. Coloration in life, dorsum, dorsal surface of head, neck, arms, and legs brown; lateral sides of dorsum and tail with many small and irregular, cream spots; venter, ventral surface of head, neck, arms, and legs dark gray, flanks have intense cream spots. Coloration in preservative as in life, but less intense. Variation. Adult male MUBI 17727 has two suboculars on the right side, males have 9-12 temporal scales and females 6-9, males are more robust than females;n paratype PFAUNA 432, the nasal suture scale does not divide the nasal, but connects inferiorly to the nostril; in MUBI 17727, the nasal suture connects inferiorly and superiorly to the nostril; and the nasal sutures of the other paratypes do not connect with the nostril and extend posteriorly to middle part of the nostril. Morphometric characters vary by sex and are shown in Table 3. Distribution and ecology. Proctopous shurugojalcapi is known only from the type locality, Area de Conservacion Privada Llamapampa-La Jalca, from 2,940—2,990 m asl (Fig. 10), District of La Jalca, Province of Chachapoyas, Department of Amazonas (Fig. 6). All specimens (eight) were found during diurnal surveys from 10:00 to 14:00 h. The new species was found in sympatry with December 2023 | Volume 17 | Number 1 | e328 Two new species of Petracola from Peru Stenocercus sp., Liophis sp., and frogs Gastrotheca monticola, Pristimantis schultei, P. cf. corrugatus, P. sp., and Centrolene sp. The habitat of P. shurugojalcapi is dominated by trees of the genera Weinmannia, Clusia, and Symplocos, and shrubs of the family Melastomataceae. Conservation. Petracola shurugojalcapi was found during field studies in support of the establishment of the Area de Conservacion Privada Llamapampa-La Jalca. The Conservation Area was established on 17 April 2015, protecting more than 26,000 ha of montane forest and wet grasslands. However, this important area 1S not exempt from threats such as deforestation and land use changes associated with local subsistence farming, ranching, and forest fires. Although available data indicate that this species could have a restricted distribution, adjacent areas remain unexplored and could host additional populations. Therefore, we recommend that P. shurugojalcapi be categorized as Data Deficient in the Red List of the International Union for Conservation of Nature (IUCN 2022). Discussion Our ML phylogenetic analysis recovered the monophyly of Petracola with respect to a clade including Cercosaura, Cercosaura manicata_ boliviana, Dendrosauridion, Proctoporus, Potamites, and Wilsonosaura, a result that is congruent with previous studies (Moravec et al. 2018; Rojas-Runjaic et al. 2021; Mamani et al. 2022). All generic lineages were recovered with high support, except Proctoporus and Pholidobolus. The genus Proctoporus has not been conclusively defined, with some studies supporting its monophyly (e.g., Goicoechea et al. 2012; Mamani et al. 2022: Sanchéz-Pacheco et al. 2018; this study), while others do not (e.g., Rojas-Runyaic et al. 2021; Torres-Carvayal et al. 2016; Vasquez-Restrepo et al. 2020). In fact, the topologies of phylogenetic trees differ depending on the methods of phylogenetic reconstruction (Bayesian inference and Maximum Likelihood; Mamani et al. 2022; Moravec et al. 2018). Sanchéz-Pacheco et al. (2018) showed that Proctoporus is monophyletic when using Maximum Parsimony approximation with either the combined data set of DNA + morphology or the dataset of DNA only. Sanchéz-Pacheco et al. (2018) also recovered the monophyly of Oreosaurus (including O. serranus). However, recent studies showed that Oreosaurus 1s not monophyletic (e.g., Mamani et al. 2022; Rojas-Runjaic et al. 2021). Despite conflicting results, progress from studies using molecular phylogenetic analyses have improved tree topologies overall. Our phylogenetic analysis recovered multiple diagnosable lineages within Petracola, including P. amazonensis, P. shurugojalcapi (UB=100), P. ventrimaculatus (UB=100), and the P waka species complex (UB=100). This preliminary result shows that the species from the right side of the DVMR are an independent lineage with respect from those of the left side (P. ventrimaculatus and P. waka). However, our sampling Amphib. Reptile Conserv. is considered as only preliminary (including only two species) and should be taken with caution. Finally, the phylogenetic position of two species that inhabit the right side of the DVMR, P. angustisoma and P. pajatensis, 1s unknown. If our hypothesis is correct, P. angustisoma and P. pajatensis should form a monophyletic lineage together with P. amazonensis and P. shurugojalcapi. Despite the low number of localities sampled, our results supported the hypothesis proposed by Kizirian et al. (2008) that P ventrimaculatus and P. waka are cryptic species complexes. Likewise, the populations of P. waka and P. ventrimaculatus are independent lineages at the species level with high genetic distance and should be recognized as such (Fig. 1, Table 2). Furthermore, we describe two of these species collected on the right side of the DVMR, which increases the diversity of the genus to seven species. The first available sequence of P waka in GenBank (named as Proctoporus sp (KU212687) by Castoe et al. 2004) was collected from Abra Gelic, a remote locality (80 km) north of the type locality (Cajabamba), and we added samples from the Cajabamba and Bafios del Inca 70 km northwest of the type locality. The molecular evidence suggests that specimens from both Abra Gelic and Bafios del Inca belong to undescribed species that will be treated in a separate study. The landscape between these locations 1s heterogeneous and includes the Huamachuquino River, which is characterized by having arid vegetation (pers. obs.) that could promote vicariance. Finally, our results suggest that the populations of Petracola onthe right side of the DVMR are evolutionarily independent from the populations on the left side. These results are similar to those proposed for birds (Hazzi et al. 2018; Weir 2009; Winger and Bates 2015), supporting the hypothesis that the DVMR promotes allopatric speciation of Andean lineages. However, more extensive sampling is needed to determine whether all Petracola species on the right side are monophyletic, and to uncover the true diversity of this rarely studied group. Acknowledgments. We thank Cesar Aguilar (MUSM), Pablo Venegas (CORBIDI), the staff of MUBI, and Evaristo Lopez (MUSA) for allowing access to their herpetological collections; and to Tiffany M. Doan and David Kizirian for reviewing our manuscript. JCC is grateful to Alexander Pari, who shared fieldwork and contributed with some lizard collections; JCC field work was funded by Deutsche Gesellschaft fur Internationale Zusammenarbeit (GIZ). The specimen collection and DNA extraction was authorized by the Servicio Nacional Forestal y de Fauna Silvestre (SERFOR), permits RDG N° 369-2019-MINAGRI-SERFOR-DGGSPFFS, RDG N° 026-2018-MINAGRI-DGGSPFFS, and RDG N° 024-2017-SERFOR/DGGSPFFS. In addition, we thank the NGO Nature and Culture International for its support in the field studies and the Rural Community of Jalca Grande for taking part in the conservation of the forests of the Area de Conservacion Privada Llamapampa-La Jalca. December 2023 | Volume 17 | Number 1 | e328 Mamani et al. Literature Cited Boulenger GA. 1900. Descriptions of new batrachians and reptiles collected by Mr. P.O. Simons in Peru. Annals Magazine of Natural History 6(7): 181-186. Castoe TA, Doan TM, Parkinson CL. 2004. 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Winger BM, Bates JM. 2015. The tempo of trait divergence in geographic isolation: avian speciation across the Marafion Valley of Peru. Evolution 69(3): 772-787. Luis Mamani is a Peruvian biologist who obtained his M.Sc. degree from the Universidad de Concepcion (UdeC) in Chile. Currently, he 1s a Ph.D. student working on the ecology, systematics, taxonomy, and evolution of gymnophthalmid lizards in the Cordillera de los Andes. Victor J. Vargas Garcia is a Biologist and Master’s student in Applied Ecology. He has more than 10 years of experience tn the study of amphibians and reptiles in the Peruvian Andes, participating and collaborating with various expeditions and scientific publications. Currently, he works on several research and conservation projects on threatened amphibians. In addition, he is a researcher associated in the Coleccion Cientifica Pro Fauna Silvestre de Ayacucho, a member of the Asociacion Herpetologica del Peru, and provides support to the IUCN Amphibian Specialist Group. He currently works at the Servicio Nacional Forestal y de Fauna Silvestre (SERFOR) tn Peru. Juan C. Chaparro is a Peruvian Biologist with extensive experience in studying the fauna of all the traditional geographic regions of Peru. Juan graduated in Biological Sciences from Universidad Nacional Pedro Ruiz Gallo, Lambayeque, Peru, and recetved a Master’s degree in Biodiversity in Tropical Areas and Conservation in 2013 from an institutional consortium of the International University of Menendez Pelayo (UIMP-Spain), Universidad Tecnolégica Indoamérica (UTIEcuador), and Consejo Superior de Investigaciones Cientificas (CSIC-Spain). He is currently the Director and Curator of the Herpetological Collection of the Museo de Biodiversidad del Peru (MUBI), and he works as a consultant in environmental studies. Juan has authored or co-authored 58 peer reviewed scientific papers, notes, book chapters, and books on fauna, especially in herpetology and arachnology, on topics such as taxonomy, biodiversity, systematics, phylogeny, conservation, and biogeography in South America. He is interested in those topics, as well as life history, distributional patterns, and evolution using amphibians and reptiles as biological models. Alessandro Catenazzi is Associate Professor at Florida International University in Miami, USA. with CORBIDI. Amphib. Reptile Conserv. He works on the systematics and conservation of Neotropical amphibians and reptiles, and the ecological dimensions of biodiversity. For his first line of research, he works primarily in the Andes and the Amazon. Alessandro collaborates with colleagues in museums and other research institutions in describing new species and developing phylogenies of amphibians and reptiles from the mega- diverse eastern slopes of the Andes. A major theme of his current research is exploring the effects of fungal diseases on the ecology of amphibians, a group that is experiencing staggering biodiversity losses worldwide. After documenting the collapse of a species-rich amphibian assemblage, he 1s interested in developing strategies to mitigate the impact of the fungal disease chytridiomycosis on the surviving species. Alessandro is an Amphibian Red List Authority and an Associated Researcher December 2023 | Volume 17 | Number 1 | e328 Mamani et al. im 100 Neusticurus_bicarinatus MRT _S6B462 Neusticurus_rudis_ MRT_526008 Placosome_cordylinum LE 1006 Kataphraktoseurus_ungerherilton MHNLS._ 1960 Pues Fchinasaura brachyceptela OCAZ_10824 Echinsaura_horrida OCAZ BEGG 100 100 Echinaseura_keyi_OCAZ_8074 99 Echinosaura_orcesi OCAZ 6299 rt = Extimsawre_palreri_MHUARIZGTI q7 Echinosaura_centralis MHUARI2387 Echinosaura_panarrensis_CHB48? in ‘00 Ancinasaura_afraria RH Andingsaura_leevis_WEI330 Andinosaura_vieta_ OCAZI0456 ‘00 _ Andinosaura_aurea_OCA7 860 Andinosaura_vespertina_OCAZ10306 ag 8 100 k Andinosaura_kiziriani_ODAZSG67 in 4 Andmosarra_crypta OCAZIO455 Andinasaura_ocuata_OCAZIO4I0 Andinosaura_hyposticta_OHMECNI 360 Centrosaura_epoderra_ MT R3S4E6 Riare_anatoloras_OCAZS169 Riarre_raneyi_OCAZ9034 Riare_coloreromen_OCA78753 Riare_sinotera_KU_217207 Riara_cashceensis OCA2 10754 Riarre_lebionis OCAZI0412 Riare_yurborum OCAZ10822 Riare_meleagris_ ODAZSB41 Riare_stigratorel_ODA27374 Riare_unicelor OCAZ 9662 Riare_calurbiana_ICNIIZ98 Riarme_balneator_ OCAZ_ 11093 Riarre_orvesi_OCAZ9035 Riare_striata MARS33 Gelanesaurus_cochranae ODA? 5587 Gelanesaurus flavoguiaris OCA2_4bll Anada_ocalata_ SMPBOD95 Aredia petersi OCAZ_50B8 Anada_rhorbiera OCAT_G873 fusponchlus_excelsum CORBIOL 14965 Macrophalidus. amectens OCAZ_ {1121 Macropholicus huencabarrbee CORGIDI_10493 Phalidohous.afiris OCAZ_94l Phalidobolus_mentium OCAZ0S6 Pholidobolus_prefrontais OCA2 9908 Pholidobolus_dicrus OCAZ 5304 Pholidobolus vertebralis OCA2 688 Pholidcbolus_mecbrydei_OCAZ_9932 Pholidobolus_tillis_ OCA2_SDO0 Pholidobolus_ulisesi_JCM_ 310 Pholidobolus_sp.|_JEM_239 Rheosdurus_sucerostrum ROM 22892 Rheosaurus_sulcarastrum ROM43B05 Oreosaurus_achlyens ENSHOIO Oreasaurus_shrevei_UWZM2U0I1.7 Orensaurus_mediarrridi_IRSNB_2674 Orecseurus_serranus_JJSS4B 100 Ceronsaura_arguus_MPEGZISB0 Cereosaura_argulus NMPEV_ 72184 Cereasaura_oshaughnessyi_LSUMZ_HI2581 Cercasaura_oshaughnessyi_NMPEV_71/60_1 Cercosaura_parkeri_LEIS60 Cercosaura_schreibersii LG 427 Cercosaura_bassleri_CORBIDI_15187 Cercosaura_ocallata MRT 977406 Cercosaura_eigenmenni_ NMPGV_ 73112 Cercosaura_eigenrann_ MPEGZI590 Cereosaura_doaee CORBI! G50 Cercnseura_pacha_MUBI 14518 Cereasaura_renicate CORBIDI 8837 Cercosaura_menicata OCA? 9793 Cercosaura_anomela MUBL 13626 Potamites_ecrlegpus NNPEV 73186 | Potarites. eepleopus CORBIDL 14382 Potarites ecolenais OCAZ_ 4599 Potarites.erythrocufaris CORBIDI 15153 Potamites_juruazensis_ CORBIDI_15579 Pctarites, rentarioola CORGID| 10791 Potarites. strangdatus OCAZ_ 6133 Potarrites_trachodus_CORBIDI_15489 Cercosaura_menicata_boliviana_ CORBIOI Sdvasaura_brava_ MUSM_32718 Selvasaura_evasa_ CORBIDI_ISI7 Selvasaura_almendarizae OCAZ 12798 Oendrosauridion_yanesha_MUSM_25345 Proctoporus_bdivianus MNCN_ 8990 Proctoporus_Cal_MHNC5322 ag Proctoporus_carebaya, MHNC5428 100 Proctoporus_kiziriani_CORBIDI_14710 547 Proctoporus_iridescens MHNCS399 a Proctaporus_lacertus_UTA_R-51487 47 a7 + = Proctoporus xestus MNCN 2425 aod Proctoporus_chasqui_MUSM_3Il72 | ig Proctoporus_spinalis MUSM_SIIBZ 14 Proctoporus_spinalis CORBIOL 7246 Proctoporus_sp._|_ CORIO! 89 | 42 Proctoporus oreades CORBIDL 7217 54 | 5g Proctoporus_pachyurus_CORBIDI_N8!! 49 Poctoporus_sp._S_MHNC&E89 too} [oa Proctoporus.ratm_MHNCII439 Proctoporus_sp._MHNC4599 Proctoporus_sucullucu_MNCN44478 98 - — Proctnnorus_Ca?_AMNH_R-IS1695 in0 Proctoporus quentheri, MUBI_|O278 Procteporus_quentheri_CORBID|_15558 ‘on Proctaporus_sp._3_CORBIDI Proctoporus_sp._4_MLUSM 80 Practaporus_katerynae_MUBI 10482 a mo Proctoparus_optimus. MUBI 2915 — Proctoporus_unsaacae_UTA_R-51488 Proctonorus, sa? CORAIDI Wiisonasaura_josyi_CORBIDI 13634 Wilsonasaura_jasyi_MUSM_3l188 (QO) Petracola_weka WUBI 26] (00 Petracola_woka MUBI 2609 100 (00) Petraccla. waka MUBL 2B05 ed Petracala_weka MUBI_2603 Petracola. weka_KU2|2687 100 100 Petracola_ventrimaculatus KU2ISB38 Petraoola_ventrimacuatus CORBIDI_ 9235 {00 Petracola_amezonensis MUBI_I1473 100 Petracda_shurugejalcapi_MUBL 17727 Petracola_shuruggjaleap_PFAUNA 430 1 ‘Alopoglsssus_atriventris MPEG 78120 al Feolengus gaudichaud_ LG. [258 i Riolara_inopinata_IRSN_2680 ________________ glare leucosticta_ IRSNBIBIS2 ‘00 100 Bechia darby MRT_S77273 Bachia_flavescens LSUMZ_HI2977 Bachia_bresslaui_MRT_SIGBB3 2B Gyrmophthedlmus_leuconystax_MRT_S4B613 Rhachisaurus_brachylenis MRT_8B7336 100 ol Fig. S1. Phylogenetic relationships of Cercosaurini (log likelihood = -54268.289, ultrafast bootstrap = 10,000) constructed from the data set of 2,384 nucleotides for mitochondrial genes (12S, 16S, cyt-b, and ND4) and a nuclear gene (c-mos). Amphib. Reptile Conserv. 173 December 2023 | Volume 17 | Number 1 | e328