Integrative revision of the Lygodactylus gutturalis (Bocage, 1873) complex unveils extensive cryptic diversity and traces its evolutionary history Author Lobón-Rovira, Javier Author Bauer, Aaron M. Author Pinto, Pedro Vaz Author Trape, Jean-Francois Author Conradie, Werner Author Kusamba, Chifundera Author Júlio, Timóteo Author Cael, Garin Author Stanley, Edward L. Author Hughes, Daniel F. Author Behangana, Mathias Author Masudi, Franck M. Author Pauwels, Olivier S. G. Author Greenbaum, Eli text Zoological Journal of the Linnean Society 2024 Zool. J. Linn. Soc. 2023-09-23 201 2 447 492 http://dx.doi.org/10.1093/zoolinnean/zlad123 journal article 301345 10.1093/zoolinnean/zlad123 ce8c1520-11a3-4d2f-b1ad-224f830e4c45 0024-4082 13302833

Table 3. Summary of external and osteological morphological data of all members within the L. gutturalis complex.Measurements are shown in mm (average and SD). For abbreviations see Material and methods section.

Species	L. gutturalis s.s.	L. dysmicus	L. depressus	L. kibera sp. nov.	L. karamoja sp. nov.	L. mirabundus sp. nov.	L. leopardinus sp. nov.	L. gamblei sp. nov.
N	(N = 27)	(N = 1)	(N = 8)	(N = 12)	(N = 6)	(N = 1)	(N = 4)	(N = 16)
Elevation (m a.s.l.)	10–761	665	7–331	811–1705	1193–1730	551	305	634–875
State of postnasal scale	81.5% not in contact	In broad contact	70% not in contact	In contact	70% not in contact	Not in contact	In contact	15% not in contact
Gular lines pattering	2–3	2	2	3	3	2 1/2	Broken pattern	2 1/2 or 3
Maximum SVL	36.2 (N = 61)	27.6	38.0	37.7	37.7	34.8	34.7	39.8
SVL	33.3 ± 1.8	27.6	35.6 ± 1.4	34.7 ± 2.5	34.4 ± 2.9	34.8	33.6 ± 1.3	36.1 ± 2.1
TL	33.6 ± 2.3	25.5	25.3 ± 0.0	37.5 ± 3.1	35.0 ± 2.8	–	32.7 ± 1.7	36.8 ± 2.6
TRL	15.3 ± 1.3	10.3	16.2 ± 1.8	14.9 ± 1.5	14.5 ± 2.6	15.4	15.0 ± 1.2	16.2 ± 1.3
HL	9.0 ± 0.6	8.3	9.2 ± 0.4	9.4 ± 0.8	9.4 ± 0.7	8.9	9.2 ± 0.5	10.1 ± 0.8
HW	6.2 ± 0.6	6.2	6.6 ± 0.2	6.7 ± 0.6	6.9 ± 0.6	7.4	6.4 ± 0.3	7.2 ± 0.6
OD	2.3 ± 0.2	2.1	2.1 ± 0.3	2.3 ± 0.2	2.1 ± 0.3	1.8	2.2 ± 0.3	2.4 ± 0.2
EarL	0.7 ± 0.1	0.7	0.8 ± 0.1	1.0 ± 0.1	0.9 ± 0.2	1.1	0.7 ± 0.1	0.8 ± 1.0
CL	5.8 ± 0.5	5.1	5.6 ± 0.6	5.7 ± 0.5	5.2 ± 0.6	6.1	5.5 ± 0.2	6.2 ± 0.8
FL	5.2 ± 0.5 (N = 24)	4.7	5.0 ± 0.5	4.7 ± 0.4	5.0 ± 0.3	5.0	4.9 ± 0.2	5.4 ± 0.3 (N = 10)
ForeL	9.5 ± 0.8	8.1	8.5 ± 2.5	9.5 ± 0.6	8.8 ± 0.4	9.6	9.4 ± 0.6	10.2 ± 0.9
NE	3.1 ± 0.3	3.1	3.5 ± 0.3	3.4 ± 0.3	3.4 ± 0.3	3.8	3.3 ± 0.3	3.6 ± 0.3
SE	4.3 ± 0.3	3.7	4.5 ± 0.3	4.6 ± 0.4	4.5 ± 0.3	4.5	4.2 ± 0.2	4.6 ± 0.4
EE	3.0 ± 0.5	2.4	2.9 ± 0.3	3.2 ± 0.4	3.3 ± 0.4	3.6	3.1 ± 0.2	3.2 ± 0.2
IN	1.8 ± 0.2	1.2	1.6 ± 0.2	1.6 ± 0.1	1.7 ± 0.3	1.8	1.6 ± 0.1	1.4 ± 0.1
IO	3.7 ± 0.3	4.0	4.0 ± 0.4	4.1 ± 0.4	3.9 ± 0.3	3.8	4.4 ± 0.5	4.1 ± 0.3
No. lamellae 4th toe	5–6	4	5	5	5	5	5	5
No. postmental scales	2–4	2	2	2–3	2–3	3	2	3
No. post-postmental	4–6	4	4–5	5–6	4–7	5	4–5	5–6
scales
No. supralabial scales	6–7/6–8	7/7	7–9/7–9	7–9/7–9	6–8/7–8	8/7	7–8/8	6–9/6–9
(right/left)
No. infralabial scales	6–7/6–8	6/7	6–7/6–7	5–7/6–7	6–7/5–7	6/6	6–7/6–7	6–7/6–7
(right/left)
No. internasal scales	1–3	2	1–2	1–2	1–2	1	2–3	1–2
No. postnasal scales	1	1	1	1	1	1	1	1
No. ventral scales	15–20	21	16–19	16–18	16–18	21	14–15	15–19
No. scales between	17–24	25	22–29	19–28	19–22	19	30–36	20–25
eyes
No. dorsal scales	–	–	–	–	–	–	–	–
across body
No. preanal pores	5–8 (x=6.73)	9	7	7–8 (x=7.17)	8	10	7	7–9 (x=7.5)
Lamellae tail tip	5–6	6	–	5–6	6	–	6	5–6
OSTEOLOGICAL	(N = 7)	(N = 1)	(N = 7)	(N = 11)	(N = 6)	(N = 1)	(N = 1)	(N = 2)
FEATURES (N)
Skull length	9.2 ± 0.6	8.2	9.5 ± 0.5	9.9 ± 0.5	9.1 ± 0.5	9.4	9.6	9.0 ± 0.8
Skull width	4.6 ± 0.4	5.0	5.5 ± 0.2	5.9 ± 0.3	5.6 ± 0.5	6.0	5.5	5.2 ± 0.3
Jaw length	8.1 ± 0.5	7.2	8.3 ± 0.4	8.6 ± 0.5	8.0 ± 0.8	7.9	8.3	7.2 ± 0.9
Premaxillary tooth loci 11		11	9–11	9–11	9–11	11	11	11
Maxillary tooth loci	16–20	16	16–18	16–19	16–18	15	18	18
(right)
Maxillary tooth loci	16–20	16	16–18	16–19	16–18	15	18	17–18
(left)
Dentary tooth loci	20	20	18–21	20–24	20–22	19	21	20–22
(left)
Frontal	Fused/semifused	Paired	Fused/ semifused	Fused/semifused	Fused	Semifused	Fused	Semifused
Parietal	Paired	Paired	Paired	Paired	Paired	Paired	Paired	Paired
Postorbitofrontal	Small/absent	Small	Small/ medium	Small/medium	Small/ medium	Small	Medium	Small/ medium
Braincase	Fused/semifused	Unfused	Fused/ semifused	Fused/unfused/	Fused/ unfused/	Unfused	Semifused	Unfused
				semifused	semifused
Compound	Fused/semifused	Unfused	Fused/unfused	Fused/ unfused/	Fused/ unfused	Unfused	Fused	Semifused
bone + surangular				semifused
Jugal	Small	Small	Absent/small/me-	Small/absent	Small	Small	Absent	Absent
			dium