Between sand, rocks and branches: an integrative taxonomic revision of Angolan Hemidactylus Goldfuss, 1820, with description of four new species Author Lobon-Rovira, Javier https://orcid.org/0000-0003-4380-9427 CIBIO Centro de Investigacao em Biodiversidade e Recurso Geneticos, Universidade do Porto, Rua Padre Armando Quintas, Campus de Vairao, 4485 - 661 Vairao, Portugal & Faculdade de Ciencias da Universidade do Porto, Porto, Portugal j.lobon.rovira@hotmail.com Author Conradie, Werner https://orcid.org/0000-0003-0805-9683 Port Elizabeth Museum (Bayworld), P. O. Box 13147, Humewood 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 Author Iglesias, David Buckley Departament de Biologia (Genetica), Facultad de Ciencias, Universidad Autonoma de Madrid (UAM), c / Darwin 2, 28049, Madrid, Spain & Centro de Investigacion en Biodiversidad y Cambio Global CIBC-UAM, Facultad de Ciencias, Universidad Autonoma de Madrid, c / Darwin 2, 28049 - Madrid, Spain Author Ernst, Raffael https://orcid.org/0000-0001-6347-1414 Museum of Zoology, Senckenberg Natural History Collections Dresden, Ko ̈ nigsbru ̈ cker Landstr. 159, D- 01109, Dresden, Germany Author Verissimo, Luis Fundacao Kissama, Rua 60 Casa 560, Lar do Patriota, Luanda, Angola Author Baptista, Ninda https://orcid.org/0000-0002-2859-6606 CIBIO Centro de Investigacao em Biodiversidade e Recurso Geneticos, Universidade do Porto, Rua Padre Armando Quintas, Campus de Vairao, 4485 - 661 Vairao, Portugal & Faculdade de Ciencias da Universidade do Porto, Porto, Portugal & TwinLab CIBIO / ISCED, Instituto Superior de Ciencias da Educacao da Huila, Rua Sarmento Rodrigues s / n, Lubango, Angola Author Pinto, Pedro Vaz CIBIO Centro de Investigacao em Biodiversidade e Recurso Geneticos, Universidade do Porto, Rua Padre Armando Quintas, Campus de Vairao, 4485 - 661 Vairao, Portugal & Fundacao Kissama, Rua 60 Casa 560, Lar do Patriota, Luanda, Angola & TwinLab CIBIO / ISCED, Instituto Superior de Ciencias da Educacao da Huila, Rua Sarmento Rodrigues s / n, Lubango, Angola text Vertebrate Zoology 2021 2021-08-09 71 465 501 http://dx.doi.org/10.3897/vz.71.e64781 journal article http://dx.doi.org/10.3897/vz.71.e64781 2625-8498-71-465 5496169A0D7D4C809B72BF0AF03A6109 06CB895DD59A53E8BF8316F91B41B0AF

Table 2. Comparison between the different Angolan Hemidactylus (excluding H. mabouia and including H. gramineus from DRC). Measurements are represented in mm (average and standard deviation), based on newly collected material for this work with exception of H. nzingae, H. gramineus and H. paivae, where morphological measurements were compiled from information available in Ceriaco et al. (2020a, 2021) and information collected for this work. For abbreviations see Material and Methods section.

Species

H. faustus sp. nov.

H. pfindaensis sp. nov.

H. carivoensis sp. nov.

H. benguellensis

H. cinganji sp. nov.

H. longicephalus LINEAGE 1

H. longicephalus LINEAGE 2

H. paivae

H. bayonii

H. vernayi

H. gramineus

H. nzingae

Specimens

n=10

n=4

n=9

n=12

n=2

n=6

n=9

n=18

n=6

n=25

n=2

n=12

SVL

39.38 +/- 3.35

45.54 +/- 3.12

38,18 +/- 5,16

49.66 +/- 8.12

48.72 +/- 5.19

46.21 +/- 8.76

47.30 +/- 3.09

57.31 +/- 2.19

36.29 +/- 3.46

32.52 +/- 4.07

36.65 +/- 4.03

44.28 +/- 3.88

TL

-

32.93

36,8 +/- 7,64

40.23 +/- 8.65

42.77 +/- 0

44.20 +/- 6.94

48.52 +/- 1.94

63.05 +/- 7.14

30.06 +/- 11.11

25.92 +/- 4.19

34.40 +/- 0.99

40.33 +/- 3.48

TRL

16.83 +/- 2.51

17.85 +/- 3.76

15,82 +/- 1,92

21.94 +/- 5.93

21.33 +/- 4.13

19.25 +/- 4.29

18.98 +/- 0.09

21.45 +/- 2.89

15.02 +/- 2.62

13.95 +/- 1.88

15.85 +/- 2.33

19.41 +/- 2.58

BW

7.44 +/- 0.93

7.43 +/- 0.41

6,99 +/- 1,92

10.056 +/- 1.83

10.14 +/- 0.50

8.28 +/- 3.26

8.5 +/- 0.81

9.81 +/- 0.47

6.69 +/- 0.78

6.12 +/- 1.26

5 +/- 0.14

7.27 +/- 1.29

FL

9.10 +/- 1.36

6.99 +/- 1.35

5,79 +/- 0,98

7.32 +/- 1.81

7.86 +/- 1.30

6.24 +/- 1.53

8.57 +/- 0.12

7.12 +/- 1.14

5.57 +/- 0.81

5.89 +/- 1.23

4.70 +/- 0.28

6.06 +/- 0.77

CL

6.32 +/- 0.54

8.03 +/- 1.73

6,75 +/- 1,18

7.74 +/- 1.47

8.39 +/- 1.98

7.11 +/- 0.76

5.47 +/- 0.92

7.21 +/- 1.47

6.30 +/- 0.63

4.90 +/- 1.18

5.55 +/- 0.21

7.03 +/- 0.68

HL

11.62 +/- 1.09

13.19 +/- 1.40

11,10 +/- 0,69

14.25 +/- 2.86

12.99 +/- 2.62

13.12 +/- 2.09

14.02 +/- 0.66

14.55 +/- 0.59

10.13 +/- 1.49

9.59 +/- 0.79

10.5 +/- 0.71

12.18 +/- 0.27

HW

8.22 +/- 0.79

8.65 +/- 0.85

6,81 +/- 1,10

9.72 +/- 1.73

9.57 +/- 0.84

8.60 +/- 1.55

9.38 +/- 0.52

10.02 +/- 0.62

6.75 +/- 0.56

6.01 +/- 0.98

6.35 +/- 0.49

8.11 +/- 0.57

HH

4.95 +/- 0.71

5.26 +/- 0.28

4,21 +/- 0,69

5.52 +/- 0.97

6.32 +/- 0.58

4.93 +/- 0.99

5.65 +/- 0.27

6.12 +/- 0.28

4.37 +/- 0.49

4.04 +/- 0.52

4.25 +/- 0.49

5.08 +/- 0.55

OD

2.87 +/- 0.29

2.81 +/- 0.18

2,68 +/- 0,36

2.97 +/- 0.39

3.14 +/- 0.13

2.97 +/- 0.48

3.02 +/- 0.11

3.21 +/- 0.18

2.42 +/- 0.14

2.25 +/- 0.29

2.10 +/- 0.42

2.81 +/- 0.34

NE

3.39 +/- 0.49

3.81 +/- 0.47

3,38 +/- 0,51

4.52 +/- 0.78

4.18 +/- 0.20

4.13 +/- 0.64

4.45 +/- 0.36

4.54 +/- 0.57

2.88 +/- 0.67

2.68 +/- 0.35

2.95 +/- 0.35

4.26 +/- 0.68

SE

4.6325 +/- 0.31

5.32 +/- 0.21

4,46 +/- 0,70

6.03 +/- 1.13

5.38 +/- 0.61

5.29 +/- 0.79

6.09 +/- 0.85

5.80 +/- 0.51

4.02 +/- 0.71

3.58 +/- 0.45

4.25 +/- 0.07

5.21 +/- 0.34

EE

3.29 +/- 0.36

3.53 +/- 0.21

2,91 +/- 0,39

3.89 +/- 0.68

3.85 +/- 0.48

3.21 +/- 0.63

3.66 +/- 0.46

3.83 +/- 0.27

2.9 +/- 0.46

2.75 +/- 0.39

2.75 +/- 0.21

3.56 +/- 0.23

EH

0.99 +/- 0.15

0.84 +/- 0.25

1,73 +/- 2,67

1.25 +/- 0.55

1.17 +/- 0.42

0.92 +/- 0.25

0.98 +/- 0.09

0.95 +/- 0.37

0.69 +/- 0.19

0.57 +/- 0.12

0.7 +/- 0.0

1.07 +/- 0.17

IN

1.56 +/- 0.24

1.36 +/- 0.05

1,21 +/- 0,20

3.75 +/- 6.35

1.42 +/- 0.12

1.41 +/- 0.21

1.59 +/- 0.15

1.63 +/- 0.18

1.27 +/- 0.12

1.22 +/- 0.18

-

-

OI

3.97 +/- 0.40

3.9 +/- 1.41

3,51 +/- 0,61

4.20 +/- 0.66

4.81 +/- 0.04

3.47 +/- 0.72

4.68 +/- 0.17

4.85 +/- 0.04

3.3 +/- 0.35

2.98 +/- 0.49

-

-

Precloacal-Femoral Pores

17-19

8

15-22

23-33

26-28

5-11

6

6-8

4-9

4-6

8

7-8

Post-cloacal (tail) spurs

1/1

1/1

1/1

1-4/1-4

1-2/1-2

0-1/0-1

1-1/1-1

-

1/1

1-2/1-2

-

-

Scales across belly

29-32

28-30

32-38

33-41

30-32

28-37

28-36

28-34

25-32

32-39

23-25

22-27

Dorsal tubercles row

15-17

11-12

12-18

13-18

15

13-17

13-16

13-16

15-16

11-16

12-14

16-21

Supralabials (right/left)

9/9

10/10

11/10-11

10-11/10-11

9/9

10-12

10-12

10-12/10-12

8-9/9-10

9-10/9-10

10/9-0

10-11/10-11

Infralabials (right/left)

8/8

8-10/9-10

9/9-10

9-10/8-10

7-9/8

9-10

9-10

9-10/8-10

7-9/7-8

7-8/7-8

8-9/8-9

8-9/8-9

Subcaudals size

Small (<1⁄4 tail width)

Small (<1⁄4 tail width)

Small (<1⁄4 tail width)

Small (<1⁄4 tail width)

Small (<1⁄4 tail width)

Small (<1⁄4 tail width)

Small (<1⁄4 tail width)

Small-Medium

Small (<1⁄4 tail width)

Small (<1⁄4 tail width)

Small (<1⁄4 tail width)

Small (<1⁄4 tail width)

# Granular scales between tubercles

2-3

4-5

2-3

2-5

2-3

3-6

3-4

3-9

2-4

2-4

2-3

2-3

Lamellae 1st finger

5-6

6-5

5

6

5-6

5-6

6

5-8

5-6

5-6

5

5-7

Lamellae 4th finger

7

7

6-7

6-8

7

7-8

7-8

8-9

7

7

7

6-8

Lamellae 1st toe

5-6

5-6

5

5-7

6

5-7

5-6

6-7

6

5-6

5

5-7

Lamellae 4th toe

9-8

7-8

9

8-9

8

8-10

8-19

9-11

8-10

7-8

7

9