Bioacoustics of Rhinolophus francisi

Echolocation Rhinolophus francisi emits a typical long narrow band FM/CF/FM signal. Based on hand-held specimens from Kalimantan and Thailand ( n = 2), the most energy is in the second harmonic with an Characters R. f. francisi X Holotype n = 3 R. f. thailandicus Y Holotype R. beddomei n = 7 R. formosae n = 42 R. trifoliatus n = 19 R. luctus n = 22 MASS – 18.0 [1] 16.00 – 20.0 ± 2.9 (11.9–25.3) 14.8 ± 1.8 (12.5–17.0) [4] 56.2 [1] FA 53.12 53.53 ± 1.01 52.90–54.70 52.97 59.83 ± 3.94 (55.00–63.44) [5] 57.58 ± 1.85 (53.85–62.40) 52.15 ± 1.90 (48.92–54.11) [10] 70.33 ± 4.44 (61.80–77.30) [17] E 25.72 25.70, 27.00 [2] 23.81 27.50, 27.60 [2] 29.32 ± 2.52 (24.45–31.30) [6] 25.78 ± 0.38 (25.35–26.20) [4] 36.11 ± 2.91 (31.48–39.60) [10] T 29.86 33.00, 38.00 [2] 30.91 37.00, 38.50 [2] 33.53 ± 2.65 (29.40–36.50) [5] 34.29 ± 1.71 (32.31–36.43) [4] 50.99 ± 6.39 (41.00–64.30) [13] TIB 27.78 29.50, 29.60 [2] 28.33 30.21 ± 1.34 28.90–31.91 [5] 31.16 ± 1.38 (28.15–33.70) [36] 26.02 ± 1.19 (24.55–28.29) [10] 36.60 ± 2.07 (33.73–40.23) [11] HF 15.20 13.50, 15.50 [2] 14.08 12.10, 13.50 [2] – 12.03 ± 0.49 (11.40–12.44) [4] 16.25 ± 2.13 (12.00–18.71) [12] NL 12.36 – 12.21 11.00 [1] – 11.23 ± 0.21 (11.00–11.43) [4] 15.25 ± 2.02 (11.70–17.15) [7] 5MET 42.85 43.36 [1] 43.99 45.28, 46.28 [2] – 43.33 ± 0.80 (42.27–44.06) [4] 58.81 ± 3.08 (55.17–62.40) [4] 4MET 41.85 41.22 [1] 42.47 44.08, 44.12 [2] – 41.39 ± 0.96 (40.22–42.39) [4] 57.05 ± 2.73 (54.14–60.09) [4] 3MET 35.94 36.15 [1] 37.47 37.77, 37.78 [2] – 36.49 ± 1.08 (35.51–38.00) [4] 50.03 ± 2.57 (47.72–53.38) [4] 3D1P 18.61 19.71 [1] 18.52 19.52, 21.55 [2] – 21.06 ± 0.84 (20.34–22.10) [4] 28.28 ± 1.02 (27.21–29.54) [4] 3D2P 26.30 26.31 [1] 27.75 27.75, 28.38 [2] – 29.20 ± 1.61 (27.74–30.93) [4] 38.99 ± 1.03 (37.96–40.32) [4] Characters R. f. francisi X Holotype n = 2 R. f. thailandicus Y Holotype R. beddomei n = 6 R. formosae n = 8 R. trifoliatus n = 14 R. luctus n = 20 R. sedulus n = 3 SL 26.57 24.27, 25.36 24.37 26.06 ± 0.96 25.55 ± 0.42 22.21 ± 0.83 30.52 ± 1.51 19.55 ± 0.59 (25.01–27.16) (24.71–26.18) (21.64–24.31) [12] (26.35–32.07) [16] (18.99–20.17) CCL 23.85 22.23, 22.75 21.72 23.41 ± 0.81 24.03 ± 0.62 20.31 ± 0.96 27.41 ± 1.01 17.42, 18.28 [2] (22.35–24.49) [5] (23.34–24.82) [5] (19.48–22.39) [11] (25.13–28.64) [16] ZB 13.51 13.00, 13.39 12.16 13.64 ± 0.75 12.67 ± 0.42 11.70 ± 0.45 15.40 ± 0.67 9.90 ± 0.39 (12.51–14.52) (11.99–13.42) (10.92–12.50) [13] (14.46–16.45) (9.67–10.35) BB 12.06 10.00, 10.51 9.93 10.82 ± 0.65 11.62 ± 0.12 9.54 ± 0.42 12.22 ± 0.42 8.25 ± 0.24 (9.80–11.54) (11.45–11.78) (8.80–10.09) [13] (11.62–13.13) (8.05–8.52) MW 12.09 11.44, 11.44 11.31 11.77 ± 0.52 10.78 ± 0.34 10.55 ± 0.30 13.34 ± 0.47 9.02, 9.63 [2] (10.97–12.25) (10.39–11.36) (10.05–10.93) [12] (12.50–13.89) [17] PC 3.00 2.28, 2.99 2.44 2.50 ± 0.33 2.47 ± 0.20 2.19 ± 0.23 3.12 ± 0.43 2.22 ± 0.14 (2.11–2.97) (2.25–2.84) (1.89–2.69) [13] (2.37–3.93) (2.11–2.38) PL 3.85 3.74 [1] 3.21 3.80 ± 0.32 3.89 ± 0.18 2.94 ± 0.26 4.55 ± 0.39 2.23 ± 0.24 (3.34–4.16) (3.68–4.19) (2.48–3.41) (3.96–5.34) (2.05–2.51) C–M3 9.78 9.73, 9.83 9.30 9.84 ± 0.48 9.88 ± 0.19 8.60 ± 0.35 11.88 ± 0.52 7.21 ± 0.24 (9.33–10.52) (9.63–10.17) (8.07–9.42) (10.62–12.81) (6.63–7.57) C1–C1 7.16 6.65, 6.91 6.07 6.71 ± 0.43 6.98 ± 0.20 8.37 ± 0.30 7.72 ± 0.43 4.63 ± 0.08 (6.09–7.24) (6.70–7.22) (7.92–8.99) (6.60–8.52) (4.55–4.70) M3–M3 9.94 9.24, 9.68 8.86 9.64 ± 0.48 9.21 ± 0.32 8.37 ± 0.30 11.02 ± 0.68 7.26 ± 0.22 (9.16–10.24) (8.78–9.55) (7.92–8.99) (9.98–12.86) (7.07–7.50) C–M3 10.88 10.13, 10.34 10.01 10.51 ± 0.51 10.42 ± 0.14 9.10 ± 0.39 12.59 ± 0.62 7.72 ± 0.53 (9.92–11.11) (10.30–10.64) (8.38–10.03) (11.19–13.84) (7.12–8.12) M 18.32 17.82, 18.11 17.15 18.34 ± 0.66 17.92 ± 0.48 15.65 ± 0.66 22.04 ± 1.10 13.41 [1] (17.46–19.13) (17.20–18.71) (14.70–16.79) [12] (19.80–23.67) AMSW 5.18 5.18, 5.18 4.76 4.86 ± 0.31 4.72 ± 0.28 4.13 ± 0.14 5.90 ± 0.31 3.98 ± 0.06 (4.46–5.18) (4.30–5.07) (3.85–4.29) (5.43–6.73) [19] (3.93–4.05) ALSW 7.35 7.02, 7.02 6.53 6.79 ± 0.40 6.77 ± 0.24 5.99 ± 0.12 8.51 ± 0.46 5.31 ± 0.16 (6.18–7.24) (6.43–7.19) (5.73–6.14) (7.32–9.06) (5.13–5.45) fmaxe of the CF segment of 49.2–50.0 kHz. The call duration ( d ) is 20.2–39.9 ms. The inter-pulse interval ( ipi ) varies between 30.7 and 126.0 ms.

Comparison with other species Rhinolophus francisi differs significantly in size from the larger R. luctus and smaller R. sedulus ( Table 1 ). It is closely similar to other medium-sized woolly horseshoe bat species, namely R. trifoliatus , R. beddomei and R. formosae , the latter two of which have a slightly longer FA ( Table 1 ) and are known to be restricted to the Indian Subcontinent and Taiwan , respectively. Although similar in size, it differs from R. trifoliatus in having brown hairs and a dark brown noseleaf and ear membranes, whereas the hairs are yellowish brown and noseleaf and the ears are yellow in R. trifoliatus . In the skull, R. francisi overlaps in size (e.g., SL and CCL) with R. beddomei and R. formosae and is slightly larger than R. trifoliatus ( Table 2 , Fig. 3 ). It is distinctly smaller than R. luctus and larger than R. sedulus ( Table 2 , Fig. 3 ). The PCA based on 13 craniodental measurements also clearly indicates the similarity of the skull size between R. francisi , R. beddomei and R. formosae and the distinction from R. luctus and R. sedulus ( Fig. 5 ). The sagittal crest of R. francisi , although well defined, is much less developed anteriorly than in R. beddomei and R. trifoliatus ( Fig. 3 ). The Taiwanese R. formosae is essentially similar in the size and shape of the skull to that of R. francisi but has distinctly shorter upper canines ( Fig. 3 ) and the anterior median swelling is slightly more inflated ( Fig. 3 ). The first upper premolar (P 2 ) of R. formosae is partially extruded so that the upper canine (C 1 ) and the second upper premolar (P 4 ) are nearly in contact ( Fig. 4c ). In R. francisi , at least in specimens examined, the P 2 is situated in the middle of the toothrow ( Fig. 4a ). In the lower toothrow, the canine (C 1 ) of R. francisi is larger than that of R. formosae in crown area ( Fig. 4a versus 4c). In R. beddomei , the teeth are more massive than in R. francisi ( Fig. 4a versus 4b). In the acoustic characters, the call frequency of R. francisi , which has an fmaxe of 49.2–50.0 kHz, is closely similar but slightly lower than that of the sympatric and similar sized species R. trifoliatus , which has an fmaxe of 50.0–53.5 kHz in Thailand ( Soisook et al. , 2010 ) and 50–54 kHz in peninsular Malaysia and Sabah ( Francis, 2008 ). Two individuals of R. beddomei recorded from Shenduruney, Agasthyamalai Hills, Kerala (southern Western Ghats), India , emitted an fmaxe of 43.0–45.5 kHz with d of 56.0–72.0 ms (K. Deshpande, unpublished data). Wordley et al. (2014) reported an fmaxe of R. beddomei as 41.7–43.3 kHz from the Anamalai Hills (southern Western Ghats), India . In Taiwan , individuals of R. formosae , which were captured and subsequently released had an fmaxe of 44.4–44.6 kHz and a d of 36.0–63.0 ms ( n = 3) (S.-F. Chen, unpublished data). The fmaxe of the larger species, R. luctus , is 32.0 kHz in Thailand ( Soisook et al. , 2010 ) and 40–42 kHz in Sabah and peninsular Malaysia ( Francis, 2008 ). The smaller species, R. sedulus , has an fmaxe of 67 kHz in peninsular Malaysia and 62–76 kHz in Sabah ( Francis, 2008 ). Character PC1 PC2 PC3 SL -0.287 0.016 0.166 CCL -0.287 0.098 0.188 ZB -0.284 0.003 -0.034 BB -0.276 0.172 0.179 MW -0.282 -0.094 0.018 PC -0.239 -0.774 -0.315 C–M3 -0.288 0.001 0.125 PL -0.273 0.259 0.267 M3–M3 -0.255 0.405 -0.835 C1–C1 -0.279 0.224 -0.064 C–M3 -0.288 -0.012 0.104 AMSW -0.276 -0.229 -0.004 ALSW -0.287 -0.138 0.078 Eigenvalues 11.772 0.447 0.225 % of total variance explained 90.550 3.440 1.730 Genetic analyses