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).

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

[_] 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.

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.

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

lat r gs

€ 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 !

: =

3 ©

ion =

® gS

3 OL Q, n

ie) in =

2. 10) 10] JO] oH ro)

o 10] )

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. Important Bird Areas

factsheet: Lake Turkana. Available at: http://www.

birdlife.org [Accessed: 1 November 2022].

Cabeza M, Fernandez-Llamazares A, Fraixedas S,

Lopez-Baucells A, Burgas D, Rocha R, Ajiko A.

2016. Winds of hope for Sibiloi1 National Park.

Swara 4: 33-37.

Ceriaco L, Marques MP, Bandeira S, Agarwal I, Stanley

EL, Bauer AM, Heinicke MP, Blackburn DC. 2018.

A new earless species of Poyntonophrynus (Anura,

Bufonidae) from the Serra da Neve Inselberg,

Namibe Province, Angola. ZooKeys 780: 109-136.

Channing A, Howell K. 2006. Amphibians of East

Africa. Comstock Publishing Associates/Cornell

University Press, Ithaca, New York, USA. 418 p. +

24 pl.

Conenna I, Lopez-Baucells A, Rocha R, Ripperger S,

Cabeza M. 2019. Movement seasonality in a desert-

dwelling bat revealed by miniature GPS loggers.

Movement Ecology 7(1): 1-10.

Cox N, Young BE, Bowles P, Fernandez M, Marin

J, Rapacciuolo G, Bohm M, Brooks T, Hedges

June 2023 | Volume 17 | Number 1/2 | e324

Kirchhof et al.

SB, Hilton-Taylor C, et al. 2022. A global reptile

assessment highlights shared conservation needs of

tetrapods. Nature 605(7909): 285-290.

Frost DR. 2022. Amphibian Species of the World: an

Online Reference. Version 6.1. American Museum

of Natural History, New York, New York, USA.

Available: https://amphibiansoftheworld.amnh.org/

index.php [Accessed: 1 April 2022].

IUCN SSC Amphibian Specialist Group. 2016.

Sclerophrys turkanae. The IUCN Red List of

Threatened Species 2016: e.T54784A 107351510.

IUCN World Heritage Outlook. 2020. Lake Turkana

National Parks - 2020 Conservation Outlook

Assessment. IUCN, Gland, Switzerland. Available:

https://rris.biopama.org/node/20655 [Accessed: 2

November 2022].

Junqueira AB, Fernandez-Llamazares A, Torrents-

Ticéd M, Haira PL, Nasak JG, Burgas D, Fraixedas

S, Cabeza M, Reyes-Garcia V. 2021. Interactions

between climate change and infrastructure projects

in changing water resources: an ethnobiological

perspective from the Daasanach, Kenya. Journal of

Ethnobiology 41(3): 331-348.

Kaiyage S, Nyagahn N. 2009. Socio-economic

Analysis and Public Consultation of Lake Turkana

Communities in Northern Kenya. Final Report.

African Development Bank, Tunis, Tunisia. 191 p.

Kirchhof S, Losogo LB, Malonza PK. 2018.

Psammophis punctulatus trivirgatus (Southern

Speckled Sand Snake). Diet. Herpetological Review

49(3): 552-553.

Largen MJ, Spawls S. 2010. The Amphibians and

Reptiles of Ethiopia and Eritrea. Frankfurt

Contributions to Natural History, Volume 38. Edition

Chimaira, Frankfurt am Main, Germany. 693 p.

Largen MJ. 2001. Catalogue of the amphibians of

Ethiopia, including a key for their identification.

Tropical Zoology 14(2): 307-402.

Liedtke HC, Muller H, Hafner J, Penner J, Gower DJ,

Mazuch T, Rodel MO, Loader SP. 2017. Terrestrial

reproduction as an adaptation to steep terrain in

African toads. Proceedings of the Royal Society B:

Biological Sciences 284(1851): 20162598.

Loveridge A. 1940. 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.

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).

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.

Elevation (m)

-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

Zambezi

River Basin

x x KK

Source of records

This study; Ruas 1996

This study; Monard 1937; Conradie

et al. 2016

This study

This study; Bocage 1895; Monard

1937; 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

This study; Monard 1937; Conradie

et al. 2016

This study; Conradie et al. 2016

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

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

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)

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:

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)

fo)

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.

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.

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)

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)

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)

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.

Elevation (m)

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=)

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.

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)

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)

Map 18. Distribution of Hyperolius aff. bocagei in Angola.

Amphib. Reptile Conserv.

Elevation (m)

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.

Elevation (m)

“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)

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

Elevation (m)

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)

oOo

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)

(=)

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.

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)

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)

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)

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)

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)

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.

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,

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)

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)

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)

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.

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. Note préliminaire sur deaux batraciens

nouveaux, des genres Rappia et Bufo, provenant

d’Afrique orientale anglaise (Mission Alluaud et

Jeannel, 1911-1912). Bulletin du Muséum National

d Histoire Naturelle. Paris 30: 269-270.

Baptista N, Conradie W, Vaz Pinto P, Branch WR.

2019. The amphibians of Angola: early studies

and the current state of knowledge. Pp 243-281 In:

Biodiversity of Angola. Science and Conservation: a

Modern Synthesis. Editors, Huntley BJ, Ferrand N,

Russo V, Lages F. Springer, Cham, Switzerland. 552

Baptista NL, Vaz Pinto P, Keates C, Edwards S, Rodel

M-O, Conradie W. 2021. A new species of red

toad, Schismaderma Smith, 1849 (Anura: Bufonidae),

from central Angola. Zootaxa 5081(3): 301-332.

Becker FS, Vaz Pinto P, Baptista N, Ernst R, Conradie

W. 2023. The amphibians of the highlands and

escarpments of Angola and Namibia. Namibian

Journal of Environment (In Press).

Benson DA, Cavanaugh M, Clark K, Karsch-Mizrachi

I, Lipman DJ, Ostell J, Sayers EW. 2013. GenBank.

Nucleic Acids Research 41(D1): D36—D42.

Bittencourt-Silva GB. 2019. Herpetological survey of

western Zambia. Amphibian & Reptile Conservation

13(2) [Special Section]: 1-28 (e181).

Bittencourt-Silva GB, Bayliss J, Conradie W. 2020.

First herpetological surveys of Mount Lico and

Mount Socone, Mozambique. Amphibian & Reptile

Conservation 14(2) [General Section]: 198-217

(e247).

Bocage JVB. 1895. Herpétologie d’Angola et du Congo.

Imprimerie Nationale, Lisbonne, France. 203 p.

Branch WR, Schmitz A, Lobon-Rovira J, Baptista NL,

Antonio T, Conradie W. 2021. Rock island melody:

a revision of the Afroedura bogerti Loveridge, 1944

group, with descriptions of four new endemic species

from Angola. Zoosystematics and Evolution 97(1):

55-82.

Branch WR, Baptista N, Vaz Pinto P, Conradie W. 2019a.

The reptiles of Angola — history, updated checklists,

endemism, hot spots, and future directions for research.

Pp. 283-326 In: Biodiversity of Angola. Science and

Conservation: a Modern Synthesis. Editors, Huntley

BJ, Ferrand N, Russo V, Lages F. Springer, Cham,

Switzerland. 552 p.

Branch WR, Conradie W, Vaz Pinto P, Tolley KA.

Amphib. Reptile Conserv.

2019b. Another Angolan Namib endemic species:

a new Nucras Gray, 1838 (Squamata: Lacertidae)

from south-western Angola. Amphibian & Reptile

Conservation 13(2) [Special Section]: 82—95 (e199).

Broadley DG. 1971. The reptiles and amphibians of

Zambia. The Puku: the Occasional Papers of the

Deptartmetn of Wildlife, Fisheries, and National

Parks, Zambia 6: 1-143.

Brooks C. 2012. Biodiversity Survey of the Upper

Angolan Catchment of the Cubango-Okavango

River Basin. USAid-Southern Africa, Pretoria,

South Africa. 151 p. Available: http://the-eis.com/

elibrary/sites/default/files/downloads/literature/

Combined%20Angolan%20Biodiversity%20

Survey%20Report_ 2012 08 05 14.pdf [Accessed: 3

April 2023].

Brooks C. 2013. Trip Report: Aquatic Biodiversity

Survey of the Lower Cuito and Cuando River Systems

in Angola. USAid-Southern Africa, Pretoria, South

Africa. 43 p. Available: http://the-eis.com/elibrary/

sites/default/files/downloads/literature/A quatic%20

biodiversity%20survey%200f%20the%20lower%20

Cuito%20and%20Cuando%20river%20systems%20

in%20Angola_2013.pdf [Accessed: 3 April 2023].

Cei JM. 1977. Chaves para uma identificacao preliminar

dos batraquios anuros da R.P. de Angola. Boletim da

Sociedade Portuguesa de Ciéncias Naturais 17: 5—26.

Ceriaco LMP, Agarwal I, Marques MP, Bauer AM.

2020a. A review of the genus Hemidactylus Goldfuss,

1820 (Squamata: Gekkonidae) from Angola, with the

description of two new species. Zootaxa 4746(1):

1-71.

Ceriaco LMP, Heinicke MP, Parker KL, Marques MP,

Bauer AM. 2020b. A review of the African Snake-

eyed Skinks (Scincidae: Panaspis) from Angola, with

the description of a new species. Zootaxa 4747(1):

77-112.

Ceriaco LMP, Agarwal I, Marques MP, Bauer AM.

2020c. A correction to a recent review of the

genus Hemidactylus Goldfuss, 1820 (Squamata:

Gekkonidae) from Angola, with the description of two

additional species. Zootaxa 4861(1): 92-106.

Ceriaco LMP, Marques MP, Bandeira S, Agarwal I,

Stanley EL, Bauer AM, Heinicke MP, Blackburn

DC. 2018. A new earless species of Poyntonophrynus

(Anura, Bufonidae) from the Serra da Neve Inselberg,

Namibe Province, Angola. ZooKeys 780: 109-136.

Ceriaco LMP, Marques MP, Bandeira SA. 2016. Anfibios

e Répteis do Parque Nacional da Cangandala.

Instituto Nacional da Biodiversidade e Areas de

Conservacao and Museu Nacional de Historia Natural

e da Ciéncia, Lisbon, Portugal. 96 p.

Ceriaco LMP, Santos BS, Marques MP, Bauer AM,

Tiutenko A. 2021. Citizen science meets specimens

in old formalin filled jars: a new species of Banded

Rubber Frog, genus Phrynomantis (Anura:

Microhylidae) from Angola. A/ytes 38(1-4): 18-48.

August 2023 | Volume 17 | Number 1 & 2 | e325

Conradie et al.

Channing A. 2022. Color patterns to sequences:

a perspective on the systematics of the

Hyperolius viridiflavus group (Anura: Hyperoliidae)

using mitochondrial DNA. Zootaxa 5134: 301-354.

Channing A. 2001. Amphibians of Central and Southern

Africa. Cornell University Press, Ithaca, New York,

USA. 470 p.

Channing A, R6del MO, Channing J. 2012. Tadpoles

of Africa. Edition Chimaira, Frankfurt am Main,

Germany. 402 p.

Channing A, Baptista N. 2013. Amietia angolensis and A.

Juscigula (Anura: Pyxicephalidae) in southern Africa:

a cold case reheated. Zootaxa 3640: 501-520.

Channing A, Dehling JM, Lotters S, Ernst R. 2016.

Species boundaries and taxonomy of the African river

frogs (Amphibia: Pyxicephalidae: Amietia). Zootaxa

4155(1): 1-76.

Channing A, Hillers A, Lotters S, Rodel MO, Schick

S, Conradie W, Ré6dder R, Mercurio V, Wagner P,

Dehling MJ, et al. 2013. Taxonomy of the super-

cryptic Hyperolius nasutus group of Long Reed Frogs

of Africa (Anura: Hyperoltidae), with descriptions of

Six hew species. Zootaxa 3620(3): 301-350.

Channing A, Rodel M-O. 2019. Field Guide to the Frogs

and Other Amphibians of Africa. Struik Nature, Cape

Town, South Africa. 408 p.

Conradie W, Baptista NL, Verburgt L, Keates C,

Harvey J, Julio T, Neef G. 2021. Contributions to

the herpetofauna of the Angolan Okavango-Cuando-

Zambezi river drainages. Part 1: Serpentes (snakes).

Amphibian & Reptile Conservation 15(2) [General

Section]: 244-278 (e292).

Conradie W, Branch WR, Measey GJ, Tolley KA. 201 2a.

Revised phylogeny of Sand Lizards (Pedioplanis)

and the description of two new species from south-

western Angola. African Journal of Herpetology

60(2): 91-112.

Conradie W, Branch WR, Measey JG, Tolley, KA. 201 2b.

A new species of Hyperolius Rapp, 1842 (Anura:

Hyperoliidae) from the Serra da Chela mountains,

southwestern Angola. Zootaxa 3269: 1-17.

Conradie W, Deepak V, Keates C, Gower DJ. 2020a.

Kissing cousins: a review of the African genus

Limnophis Gunther, 1865 (Colubridae: Natricinae),

with the description of a new species from north-

eastern Angola. African Journal of Herpetology

69(1): 79-107.

Conradie W, Keates C, Lobon-Rovira J, Vaz Pinto P,

Verburgt L, Baptista NL, Harvey J, Julio T. 2020b. New

insights into the taxonomic status, distribution, and

natural history of De Witte’s Clicking Frog (Kassinula

wittei Laurent, 1940). African Zoology 55(4): 311-

B22.

Conradie W, Keates C, Baptista NL, Lobon-Rovira

J. 2022a. Taxonomical review of Prosymna

angolensis Boulenger, 1915 (Elapoidea, Prosymnidae)

with the description of two new species. ZooKeys

Amphib. Reptile Conserv.

1121: 97-143.

Conradie W, Schmitz A, Lobén-Rovira J, Becker FS,

Vaz Pinto P, Hauptfleisch ML. 2022b. Rock island

melody remastered: two new species in the Afroedura

bogerti Loveridge, 1944 group from Angola and

Namibia. Zoosystematics and Evolution 98(2): 435—

453.

Conradie W, Branch WR, Tolley KA. 2013. Fifty Shades

of Grey: giving color to the poorly known Angolan

Ash Reed Frog (Hyperoliidae: Hyperolius cinereus),

with the description on a new species. Zootaxa

3635(3): 201-223.

Conradie W, Keates C, Verburgt L, Baptista NL, Harvey

J, Julio T, Neef G. 2022c. Contributions to the

herpetofauna of the Angolan Okavango-Cuando-

Zambezi River drainages. Part 2: Lizards (Sauria),

chelonians, and crocodiles. Amphibian & Reptile

Conservation 16(2) [General Section]: 181-214

(e322).

Conradie W, Bills R. 2017. Wannabe ranid: notes on the

morphology and natural history of the Lemaire’s Toad

(Bufonidae: Sclerophrys lemairii). Salamandra 53(3):

439-444.

Conradie W, Bill R, Branch WR. 2016. The herpetofauna

of the Cubango, Cuito, and lower Cuando river

catchments of south-eastern Angola. Amphibian &

Reptile Conservation 10(2) [Special Section]: 6-36

(e126).

Du Preez L, Carruthers V. 2017. Frogs of Southern

Africa: a Complete Guide. Struik Publishers, Cape

Town, South Africa. 520 p.

Ernst R, Lautenschlager T, Branquima MF, Holting M.

2020. At the edge of extinction: a first herpetological

assessment of the proposed Serra do Pingano

Rainforest National Park in Uige Province, northern

Angola. Zoosystematics and Evolution 96: 237-262.

Ernst R, Nienguesso ABT, Lautenschlager T, Bare} MF,

Schmitz A, Holting M. 2014. Relicts of a forested

past: southernmost distribution of the Hairy Frog

genus Trichobatrachus Boulenger, 1900 (Anura:

Arthroleptidae) in the Serra do Pingano region of

Angola with comments on its taxonomic status.

Zootaxa 3779(2): 297-300.

Ernst R, Schmitz A, Wagner P, Branquima MF, Holting

M. 2015. A window to Central African forest history:

distribution of the Xenopus fraseri subgroup south

of the Congo Basin, including a first country record

of Xenopus andrei from Angola. Salamandra 52(1):

147-155.

Frost R. 2023. Amphibian Species of the World: an

Online Reference. Version 6.1, American Museum of

Natural History, New York, USA. Available: https://

amphibiansoftheworld.amnh.org/index. php

[Accessed: 3 April 2023].

Furman BL, Bewick AJ, Harrison TL, Greenbaum E,

Gvozdik V, Kusamba C, Evans BJ. 2015. Pan-African

phylogeography of a model organism, the African

August 2023 | Volume 17 | Number 1 & 2 | e325

Amphibians of the Okavango Delta headwater area in Angola

Clawed Frog Xenopus laevis. Molecolur Ecology

24(4): 909-925.

Hallermann J, Ceriaco LMP, Schmitz A, Ernst R,

Conradie W, Verburgt L, Marques MP, Bauer AM.

2020. A review of the Angolan House Snakes, genus

Boaedon Dumeéril, Bibron, and Dumeéril (1854)

(Serpentes: Lamprophiidae), with description of three

new species in the Boaedon fuliginosus (Boie, 1827)

species complex. African Journal of Herpetology

69(1): 29-78.

Hubler N. 2015. Diversity and phylogeopraphy of

Ridged Frogs (Anura: Ptychadena) in Northern

Angola: morphological and molecular evidence.

Master’s thesis, University of Greifswald, Greifswald,

Germany.

Jongsma GFM, Barej) ME, Barratt CD, Burger

M, Conradie W, Ernst R, Greenbaum E, Hirschfeld

M, Leaché AD, Penner J, et al. 2018. Diversity and

biogeography of frogs in the genus Amnirana (Anura:

Ranidae) across sub-Saharan Africa. Molecular

Phylogenetics and Evolution 120: 274-285.

Larson TR, Castro D, Behangana M, Greenbaum

E. 2016. Evolutionary history of the river frog

genus Amietia (Anura: Pyxicephalidae) reveals

extensive diversification in Central African highlands.

Molecular Phylogenetics and Evolution 99: 168-181.

Laurent RF. 1972. Tentative revision of the genus Hemisus.

Annales du Musée Royal de |’Afrique Centrale. Série

in Octavo, Science Zoologique, Tervuren 194: 1-67.

Laurent RF. 1964. Reptiles et amphibiens de I’Angola

(troisieme contribution). Publicagées Culturais da

Companhia de Diamantes de Angola 67: 11-165.

Lobon-Rovira J, Conradie W, Baptista NL, Vaz Pinto

P. 2022. A new species of Feather-tailed Leaf-toed

Gecko, Kolekanos Heinicke, Daza, Greenbaum,

Jackman, and Bauer, 2014 (Squamata, Gekkonidae)

from the poorly explored savannah of western Angola.

ZooKeys 1127: 91-116.

Lobon-Rovira J, Conradie W, Buckley Iglesias D, Ernst R,

Verisimmo L, Baptista N, Vaz Pinto P. 2021. Between

sand, rocks, and branches: an integrative taxonomic

revision of Angolan Hemidactylus Goldfuss, 1820,

with description of four new species. Vertebrate

Zoology 71: 465-501.

Loveridge A. 1953. Zoological results ofa fifth expedition

to East Africa. 1V. Amphibians from Nyasaland and

Tete. Bulletin of the Museum of Comparative Zoology

110: 325-406.

Marques MP, Parrinha D, Santo BS, Bandeira S, Butler

BO, Sousa CAN, Bauer AM, Wagner P. 2022a.

All in all it’s just another branch in the tree: a new

species of Acanthocercus Fitzinger, 1843 (Squamata:

Agamidae), from Angola. Zootaxa 5099(2): 221-243.

Marques MP, Ceriaco LMP, Heinicke MP, Chehouri RM,

Conradie W, Tolley KA, Bauer A. 2022b. The Angolan

Bushveld Lizards, genus Heliobolus Fitzinger, 1843

(Squamata: Lacertidae): integrative taxonomy and the

Amphib. Reptile Conserv.

description of two new species. Vertebrate Zoology

72: 745-769.

Marques MP, Parrhina D, Ceriaco LMP, Brennan IG,

Heinicke MP, Bauer AM. 2023. A new species of

thick-toed gecko (Pachydactylus) from Serra da Neve

and surrounding rocky areas of southwestern Angola.

Vertebrate Zoology 73: 325-343.

Marques MP, Ceriaco LMP, Bandeira S, Pauwels OSG,

Bauer AM. 2019a. Description of a new Long-tailed

Skink (Scincidae: Trachylepis) from Angola and the

Democratic Republic of the Congo. Zootaxa 4568 (1):

51-68.

Marques MP, Ceriaco LMP, Blackburn DC, Bauer AM.

2018. Diversity and distribution of the amphibians

and terrestrial reptiles of Angola: atlas of historical

and bibliographic records (1840-2017). Proceedings

of the California Academy of Sciences Series 4

65(Supplement IT): 1-501.

Marques MP, Ceriaco LMP, Buehler MD, Bandeira

SA, Janota JM, Bauer AM. 2020. A revision of the

Dwarf Geckos, genus Lygodactylus (Squamata:

Gekkonidae), from Angola, with the description of

three new species. Zootaxa 4853(3): 301-352.

Marques MP, Ceriaco LMP, Stanley EL, Bandeira SA,

Agarwal I, Bauer AM. 2019b. A new species of

Girdled Lizard (Squamata: Cordylidae) from the Serra

da Neve Inselberg, Namibe Province, southwestern

Angola. Zootaxa 4668(4): 503-524.

Mertens R. 1938. Amphibien und Reptilien aus Angola

gesmmelt von W. Shack. Senckenbergiana 20(6):

425-443,

Monard A. 1937. Contribution a la batrachologie

d’Angola. Bulletin de la Société Neuchételoise des

Sciences Naturelles 62: 5—59.

National Geographic Okavango Wilderness Project.

2017. Initial Findings from Exploration of the Upper

Catchments of the Cuito, Cuanavale, and Cuando

Rivers, May 2015 to December 2016. Wild Bird Trust,

Johannesburg, South Africa. Available: http://www.

wildbirdtrust.com/wp-content/uploads/2018/03/

NGOWP._report_1 FOR PRINT_V14_ English. pdf

[Accessed: 3 April 2023].

Ne€as T, Kielgast J, Nagy ZT, Chifundera ZK, Gvozdik

V. 2022. Systematic position of the Clicking Frog

(Kassinula Laurent, 1940), the problem of chimeric

sequences, and the revised classification of the

family Hyperoliidae. Molecular Phylogenetics and

Evolution 174: 1-11 (107514).

Nielsen SV, Conradie W, Ceriaco LMP, Bauer AM,

Heinicke MP, Stanley EL, Blackburn DC. 2020. A

new species of Rain Frog (Brevicipitidae, Breviceps)

endemic to Angola. ZooKeys 979: 133-160.

Parrinha D, Marques MP, Heinicke MP, Khalid F, Parker

KL, Tolley KA, Childers JL, Conradie W, Bauer

AM, Ceriaco LMP. 2021. A revision of Angolan

species in the genus Pedioplanis Fitzinger (Squamata:

Lacertidae), with the description of a new species.

August 2023 | Volume 17 | Number 1 & 2 | e325

Conradie et al.

Zootaxa 5032(1): 146.

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).

[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.

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.

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.

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

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.

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.

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,

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 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.

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.

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.

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.

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

5 4. 9.5 11.2 13.6 14.3 13.3

California (12.2) (18.3)

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

(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

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

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

vay

[oe

| Q

‘a beara

“N

+ | al

Ne)

ia)

—

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

| 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

+])+]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

| 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+

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.

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.

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

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:

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

Zlzazilzizlz\lz\|Z

ich

18)

NE3

13)

NE3

FE.

NE3

Environmental Vulnerability

Category (Score)

———

————

_———

—— |

——<——=——

IUCN

categorization

LG,

ill

RG,

status

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

[ay Odd

a a

——

a i

Sauromalus ater | NEB.

Sauromalus hispidus** Ea PES |

Sauromalus klauberi** | PECd

Sauromalus slevini** | PECd

[es

rey

[rey

Phrynosona blaimilti | NES Ss M(2)——SS—S

—) a

_——

| NES

| NE

|e PE een ©

a a

—_—

| NES

Callisaurus draconoides C

Petrosaurus mearnsi s

Petrosaurus repens**

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)

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 or:

Uta lowei™ rey

| NES

La a

| NES

LL Eee |

[-_*PEe_s|

| NES

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**

YE3

Plestiodon gilberti ES

iG,

TAS

LG.

KG,

LG,

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

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**

Aspidoscelis pictus**

Aspidoscelis tigris

Xantusia gilberti**

Xantusia henshawi

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’

Le.

Pituophis insulanus** FP LG

Pituophis vertebralis** P Ee

Rhinocheilus etheridgei** ‘ig

Rhinocheilus lecontei Le

Salvadora hexalepis

Sonora annulata

Sonora cincta**

Sonora fasciata**

Sonora mosaueri** E

™

Sonora punctatissima**

Amphib. Reptile Conserv. 87 November 2023 | Volume 17 | Number 1 & 2 | e326