A taxonomic revision of the whitefish of lakes Brienz and Thun, Switzerland, with descriptions of four new species (Teleostei, Coregonidae) Author Selz, Oliver M. Department of Fish Ecology and Evolution, Centre for Ecology, Evolution & Biogeochemistry, Eawag: Swiss Federal Institute of Aquatic Science and Technology, 6047 Kastanienbaum, Switzerland & Aquatic Ecology and Evolution, Institute of Ecology and Evolution, University of Bern, 3012 Bern, Switzerland https://orcid.org/0000-0002-2210-5909 oliver.selz@eawag.ch Author Doenz, Carmela J. Department of Fish Ecology and Evolution, Centre for Ecology, Evolution & Biogeochemistry, Eawag: Swiss Federal Institute of Aquatic Science and Technology, 6047 Kastanienbaum, Switzerland & Aquatic Ecology and Evolution, Institute of Ecology and Evolution, University of Bern, 3012 Bern, Switzerland Author Vonlanthen, Pascal Department of Fish Ecology and Evolution, Centre for Ecology, Evolution & Biogeochemistry, Eawag: Swiss Federal Institute of Aquatic Science and Technology, 6047 Kastanienbaum, Switzerland & Aquabios GmbH, Les Fermes 57, 1792 Cordast, Switzerland Author Seehausen, Ole Department of Fish Ecology and Evolution, Centre for Ecology, Evolution & Biogeochemistry, Eawag: Swiss Federal Institute of Aquatic Science and Technology, 6047 Kastanienbaum, Switzerland & Aquatic Ecology and Evolution, Institute of Ecology and Evolution, University of Bern, 3012 Bern, Switzerland text ZooKeys 2020 989 79 162 http://dx.doi.org/10.3897/zookeys.989.32822 journal article http://dx.doi.org/10.3897/zookeys.989.32822 1313-2970-989-79 F78F6D879DDB4CD98E4C60E4883A59B6 65AB71A5B3985F5E9FBF8CD56BC96143

Table 11. The first- and second-best ratios retrieved from the LDA ratio extractor of either head or body characters (see Table 1) alone or combined, used for pair-wise comparisons of all contemporary specimens from the four whitefish species of Lake Brienz. For some species comparisons only a subset of characters could be used (a-l); the respective characters that were excluded are listed at the end of the table. Only external characters were used for the LDA comparisons, since internal characters (gill raker and gill arch length) cannot be measured on live specimens, and are thus not informative to assign specimens to species in the field. Due to large size differences between the species the LDA ratios were calculated with three different datasets; once each with individuals larger or smaller than 163.5mm standard length and once with the full size ranges of all species. For the multi-species comparisons, only the comparisons that yielded distinguishing ratios are shown. δ is a measure of how good shape discriminates in comparison to size (i.e., the smaller the less allometry). Ratios marked with an asterisk * have very little (for the pairwise species comparisons not more than one specimen of one species overlaps with that of the other species) or no overlap and were thus eligible for use in the species key and the diagnoses.

Characters	Species comparison	Size range	Best ratios	Range species 1	Range species 2	Standard distance	δ (Shape vs. size)
(a) PelvS, PecF1, DFAd, DFAe, DFPe, TL, SL, EH, SD, SW, INW, IOW (b) PelvFS, PelvFB, PecFB, DFPe, TL, EH, ED, SD, IOW (c) PelvFB, PelvFS, PelvF, PecF1, DFB, DFAe, DFPe, AFB, AFAe, AdFB, CF, PAdC, PreP, PreA, SL, TL, BD, PostD, DHL, ED, EH, ES, PostO, HD, MW, SN, SD, SW, IOW, INW (d) PecFB, PelvFB, PelvF, PelvFS, PecF1, DFAe, DFPe, AFAe, AdFB, PreP, CF, TL, PostD (e) ED, EH, HD, SD, SW, INW (f) PelvFB, PelvFS, PelvF, PecFB, DFAe, DFAd, DFPe, CF, PreP, SL, TL, ED, EH, MW, SD, SW, IOW, INW, ES (g) PelvFS, PecF1, PecFB, DFAd, DFPe, SL, TL, ED, EH, INW, CF (h) PelvS, PecF1, DFAd, DFAe, DFPe, TL, SL, EH, SD, SW, INW, IOW (i) PelvFS (j) PelvFB, PelvFS, PelvF, PecF1,DFAe, DFPe, AFB, AFAe,CF, SL, TL, BD, ED, EH, ES, PostO, HD, MW, SN, SD, SW, IOW, INW (k) PelvFS, PecF1, CF, SL, TL (l) PelvS, TL, INW
head + body	C. albellus vs. C. alpinus (a)	<163.5mm	1: PreA/LJ *	6.33-7.44	9.24-9.97	27.13	0.04
			2: AFAe/M	1.65-2.25	2.58-2.63	25.94	0.04
body	C. albellus vs. C. alpinus	<163.5mm	1: PecF2/DFAd *	0.81-1.06	0.78-0.8	9.97	0.14
			2: DHL/PreD	0.34-0.42	0.32-0.34	9.4	0.15
head	C. albellus vs. C. alpinus	<163.5mm	1: HD/LJ	1.30-1.55	1.77-1.92	15.43	0.02
			2: IOW/UJW	0.89-1.30	1.20-1.26	14.14	0.02
head + body	C. albellus vs. C. fatioi	<163.5mm	1: PecF2/PreA *	0.22-0.28	0.2-0.22	5.78	0.16
			2: DHL/PreP	0.31-0.38	0.30-0.32	4.49	0.2
body	C. albellus vs. C. fatioi	<163.5mm	1: PecF2/PreA *	0.22-0.28	0.2-0.22	6.76	0.17
			2: DHL/TL	0.13-0.18	0.13-0.14	5.7	0.19
head	C. albellus vs. C. fatioi	<163.5mm	1: UJ/ES *	6.81-12.42	4.51-6.15	8.63	0.12
			2: EH/HL *	0.27-0.31	0.23-0.27	7.3	0.14
head + body	C. albellus vs. C. brienzii (b)	<163.5mm	1: PreD/LJ *	3.99-4.68	5.05-5.57	47.9	0.01
			2: M/ES *	5.35-9.76	3.31-4.37	47.63	0.01
body	C. albellus vs. C. brienzii	<163.5mm	1: PecF2/PreD *	0.36-0.45	0.29-0.32	15.95	0.06
			2: DHL/TL	0.13-0.18	0.13-0.14	9.91	0.05
head	C. albellus vs. C. brienzii	<163.5mm	1: LJ/ES *	9.62-17.28	6.01-6.49	12.51	0.05
			2: HL/UJ	2.87-3.5	3.19-3.6	8.87	0.04
Characters	Species comparison	Size range	Best ratios	Range species 1	Range species 2	Standard distance	δ (Shape vs. size)
head + body	C. alpinus vs. C. fatioi (b)	>163.5mm	1: AFAe/UJ *	1.96-2.5	1.66-1.96	26.08	0.04
			2: CL/PreA	0.14-0.18	0.17-0.21	26.46	0.04
body	C. alpinus vs. C. fatioi	>163.5mm	1: AFae/TL	0.1-0.11	0.09-0.1	13.41	0.11
			2: CL/PreA	0.14-0.18	0.17-0.21	13.41	0.11
head	C. alpinus vs. C. fatioi	>163.5mm	1: HL/UJ *	3.55-3.93	3.13-3.55	11.51	0.07
			2: LJW/UJW	0.33-0.44	0.38-0.55	11.02	0.07
head + body	C. alpinus vs. C. brienzii (c)	>163.5mm	1: CD/SW *	2.25-2.64	1.82-2.04	34.25	0.02
			2: LJW/UJW *	0.33-0.44	0.45-0.55	33.91	0.02
body	C. alpinus vs. C. brienzii (d)	>163.5mm	1: DFAe/PAdC *	1.11-1.32	0.96-1.16	18.53	0.07
			2: CD/AFB	0.61-0.68	0.52-0.62	18.31	0.07
head	C. alpinus vs. C. brienzii (e)	>163.5mm	1: LJW/UJW *	0.33-0.44	0.45-0.55	7.44	0.08
			2: PostO/UJ	1.8-2.12	1.57-1.86	6.78	0.08
Characters	Species comparison	Size range	Best ratios	Range species 1	Range species 2	Standard distance	δ (Shape vs. size)
head + body	C. fatioi vs. C. brienzii (f)	>163.5mm	1: CL/PAdC	0.71-0.86	0.66-0.76	7.46	0.08
			2: BD/LJ	2.44-3.05	2.31-2.82	7.36	0.08
body	C. fatioi vs. C. brienzii	>163.5mm	1: CL/PAdC	0.71-0.86	0.66-0.76	6.04	0.1
			2: CF/BD	0.87-1.13	0.93-1.12	5.92	0.1
head	C. fatioi vs. C. brienzii	>163.5mm	1: ED/M	1.03-1.19	1.04-1.57	3.58	0.18
			2: HW/UJW	1.89-2.23	1.88-2.33	3.38	0.19
Characters	Species comparison	Size range	Best ratios	Range species 1	Range species 2	Standard distance	δ (Shape vs. size)
head + body	C. alpinus vs. other 3 species	<163.5mm	1: DFAd/LJ *	2.57-2.58	1.6-2.1	23.47	0.03
			2: AdFB/PAdC	0.26-0.28	0.21-0.42	22.66	0.03
head + body	C. albellus vs. other 3 species	<163.5mm	1: PostD/EH *	5.47-6.93	7.5-8.9	48.36	0.02
			2: UJW/ES	4.88-9.3	3.41-5.31	48.13	0.02
head + body	C. alpinus vs. C. fatioi + C. brienzii (g)	>163.5mm	1: DFAe/UJ *	3.28-4.1	2.58-3.19	24.71	0.05
			2: CD/SW *	2.25-2.64	1.76-2.27	24.37	0.05
Characters	Species comparison	Size range	Best ratios	Range species 1	Range species 2	Standard distance	δ (Shape vs. size)
head + body	C. albellus vs. C. alpinus (h)	100-290	1: PreD/LJ *	3.99-4.68	5.6-6.81	22.86	0.13
			2: DFAe/UJ *	2.14-2.79	3.25-4.1	21.65	0.14
head	C. albellus vs. C. alpinus	100-290	1: HD/UJ *	1.87-2.2	2.38-2.78	14.39	0.18
			2: LJ/IOW *	1.53-1.99	1.33-1.57	13.25	0.19
head + body	C. albellus vs. C. fatioi	100-290	1: PreP/EH *	6.56-7.98	8.94-11.43	15.95	0.13
			2: CL/UJ	1.44-2.02	1.93-2.72	15.09	0.14
head + body	C. albellus vs. C. brienzii (i)	100-290	1: PreD/EH *	6.1-7.58	8.12-10.32	50.86	0.04
			2: CL/LJ	0.99-1.45	1.38-1.65	50.6	0.04
head	C. albellus vs. C. brienzii	100-290	1: EH/HL *	0.27-0.31	0.22-0.27	9.33	0.18
			2: LJ/ES	9.62-17.28	6.08-12.43	8.57	0.22
Characters	Species comparison	Size range	Best ratios	Range species 1	Range species 2	Standard distance	δ (Shape vs. size)
head + body	C. alpinus vs. C. brienzii (j)	100-290	1: DFAd/LJW *	9.84-14.82	6.05-8.91	20.72	0.03
			2: DHL/LJ *	1.84-2.22	1.63-1.82	20.47	0.02
body	C. alpinus vs. C. brienzii (k)	100-290	1: PecF2/DFAd *	0.74-0.85	0.85-1.03	87.52	<0.01
			2: CD/PostD	0.17-0.2	0.15-0.18	87.48	<0.01
head	C. alpinus vs. C. brienzii	100-290	1: HD/LJW *	6.72-9.39	5.23-6.66	11.94	0.04
			2: HL/LJ *	2.54-2.96	2.19-2.47	11.61	0.04
head + body	C. alpinus vs. C. fatioi (h)	100-290	1: DFAe/UJ *	3.25-4.1	2.45-3.17	18.98	0.03
			2: PecF2/AFAe	1.24-1.47	1.37-1.63	18.63	0.03
body	C. alpinus vs. C. fatioi	100-290	1: PecF2/DFAe	0.77-0.89	0.87-1.02	9.25	0.08
			2: AFAe/PostD	0.27-0.32	0.22-0.29	8.71	0.08
head	C. alpinus vs. C. fatioi	100-290	1: LJW/UJW	0.33-0.47	0.37-0.55	5.62	0.08
			2: HL/UJ	3.43-3.93	3.13-3.63	4.98	0.08
Characters	Species comparison	Size range	Best ratios	Range species 1	Range species 2	Standard distance	δ (Shape vs. size)
head + body	C. fatioi vs. C. brienzii (l)	100-290	1: AFB/BD	0.45-0.67	0.44-0.58	28.19	<0.01
			2: PreD/M	8.87-14.85	9.13-11.41	28.16	<0.01
body	C. fatioi vs. C. brienzii	100-290	1: AFB/BD	0.45-0.67	0.44-0.58	2.76	0.05
			2: PreP/PreA	0.58-0.65	0.57-0.64	2.51	0.05
head	C. fatioi vs. C. brienzii	100-290	1: ED/M	1.04-1.57	1.03-1.28	1.72	0.1
			2: SN/MW	2.09-2.63	1.78-2.87	1.52	0.1
Characters	Species comparison	Size range	Best ratios	Range species 1	Range species 2	Standard distance	δ (Shape vs. size)
head + body	C. albellus vs. other 3 species	100-290	1: PreD/EH *	6.1-7.58	8.12-10.5	10.89	0.16
			2: CL/UJ	1.44-2.02	1.85-2.72	9.79	0.17
head + body	C. alpinus vs. other 3 species	100-290	1: DFAe/UJ *	3.25-4.1	2.14-3.19	9.59	0.11
			2: LJW/UJW	0.33-0.47	0.34-0.55	8.98	0.12