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Additional file 1: Additional figures and tables related to the statistical analyses.

Renner, Swen C. (2017): A new species of Murina (Chiroptera: Vespertilionidae) from sub-Himalayan forests of northern Myanmar. Zootaxa 4320 (1): 159-172, DOI: https://doi.org/10.11646/zootaxa.4320.1.9

Murina hkakaboraziensis sp. nov. (Figs. 1–4; Table 1) Holotype. Adult male, field number PS160218.6, (to be subsequently deposited at the Zoological Collection of the University of Mandalay (UMZC), Myanmar), body in alcohol, skull and baculum extracted. Type locality. Myanmar, Kachin, Putao Township, Hkakabo Razi National Park (proposed southern extension), 6 km north of Mali Raing Village, 27°37’19” N, 97°22’13” E, 510 m a.s.l., collected on 18 February 2016, by Pipat Soisook, Sai Sein Lin Oo and Awatsaya Pimsai. Diagnosis. A small tube-nosed bat with a FA of 29.6 mm. The external appearance is characterised by generally orange-brown pelage on the dorsal and whitish-grey on the ventral side; with long bright golden tip guard hairs on the dorsal side, most densely on the top of the head and the forehead. In the skull, the forehead profile rises smoothly to the braincase, which is relatively high but not dome-shaped. The sagittal crest and lambdoid crest are absent. The upper canine exceeds the posterior upper premolar in height and is slightly more than half of its crown area. The mesostyles on the first and second upper molars are well developed. The talonids of both first and second lower molars are equal to that of their respective trigonids in size. Etymology. The species is named after the Hkakabo Razi Landscape, where the only known specimen was collected. The proposed English name is ‘Hkakabo Razi Tube-nosed Bat’. Description. This is a very small vespertilionids with a FA of 29.6 mm and HB of 35.5 mm (Table 1). The pinna is rounded, with 17.0 mm in height, and is without distinct emargination on the posterior border. The tragus is relatively short, shorter than half of the pinna (Fig. 1a), with a height of 7.5 mm (Table 1). The body is covered with long hairs, particularly on the head (Fig. 1a). The most distinctive feature of the pelage is long bright golden hairs over the back of the head (Fig. 1a). The face and forehead are covered with very short dark brown hairs. The pelage on the dorsal side is greyish-brown basally and orange-brown at the tips (Fig. 1b); the very long and bright golden guard hairs are sparsely distributed over the back and tail membrane, but are more plentiful on the head (Fig. 1a). The pelage on the ventral side is very dark at the base, followed by grey in the middle and with a whitishgrey tip (Fig. 1c). The wing membrane is attached to the base of the claw of the outer toe. The foot is relatively large, 8.6 mm, which is more than half of the tibia length (Table 1). In the skull, the small and shortened first upper premolar (P2) indicates that this species belongs to the suillia - type. The GTL and CCL are 13.82 mm and 12.35 mm, respectively (Table 2). The rostrum is deep and bulbous. The interorbital region is deeply concave when viewed from above. The lateral profile of the forehead rises gradually and smoothly to the braincase (Fig. 2a). The braincase is relatively high (BH 6.02 mm) but not domed in shape. The sagittal crest and lambdoid crest are absent (Fig. 2a). Each zygoma is thin and without a dorsal process. The upper toothrows converge anteriorly (Fig. 2a); C1–C1 is 62.6% of M3–M3. The C–M3 is 4.75 mm (Table 2). The inner upper incisor (I2) is slightly longer than the outer one (I3) and is about half the crown area (Fig. 2a). The secondary cusp of the I3 is well developed and is about half the height of the main cusp. The I3 is relatively large, its crown area is about two-thirds that of the upper canine (C1) and subequal to that of the posterior upper premolar (P4) (Fig. 2a). The C1 is rounded in shape and without a lingual cusp at its base. It is about two-thirds that of P 4 in crown area. In the lateral view, the C1 slightly exceeds the P 4 in height (Fig. 2a). The anterior upper premolar (P2) is small, broader than long. It is slightly less than half that of the P 4 in crown area and about half the height (Fig. 2a). The labial surface of the first (M1) and second (M2) upper molars is W-shaped (Fig. 2a). The mesostyles of both M1 and M2 are well developed. ……continued on the next page In the mandible, the toothrow length (C-M3) is 5.15 mm. The lower canine (C1) is relatively small, but exceeds the anterior (P2) and the posterior lower premolar (P4) in both height and crown area (Fig. 2a). The cingulum of the C1 is well-defined and possesses a small anterior cusp. The anterior lower premolar (P2) is slightly more than half that of the C1 and about two-third that of the P 4 in height. Its crown area is about half that of the C1 and two-thirds of the P4 (Fig. 2a). The P4 is about two-thirds that of the C 1 in both height and crown area. The first and second molars (M1 and M2) are relatively large in comparison with the C1, such that the C1 is only about slightly more than half the crown area of the M1 and M2. The crown area of the talonid of both M1 and M2 is about equal to that of its respective trigonid (Fig. 2a). The coronoid process is projected slightly forward and relatively low with a CPH of 2.95 mm (Table 2). The baculum is very small with a total length of 0.8 mm and 0.6 mm in width (Fig. 3). It is somewhat rectangular in shape and without constriction (Fig. 3). The dorsal side is arch upward. Echolocation. Currently, for M. hkakaboraziensis sp. nov., echolocation calls are only known from the holotype. Unfortunately, the calls were relatively faint and it was not possible to record a good signal of the initial hook. This made it impossible to measure the start frequency (sf) accurately. Nevertheless, it is possible to describe that this Murina emits broadband frequency modulated (FM) signals typical of the genus with a highest frequency (hf) of 164.0–169.0 kHz. The energy is distributed almost evenly throughout the pulse, with an fmaxe between 148.5 and 152.8 kHz. The terminal frequency (tf) varied from 62.0–70.0 kHz. The call duration (d) was 3.0–3.8 ms and the inter-pulses interval (ipi) was 69.1–75.3 ms. Echolocation characters indicate that this species, as many other Murina, is a forest-adapted species. However, based on the habitat of the capture site (see below), it is likely that it is adaptable to forage around edges between forest and open space. Genetics. The Bayesian analysis phylogram based on DNA barcode reveals that several species in the subfamily Murininae are paraphyletic (Fig. 4). The single specimen of M. hkakaboraziensis sp. nov. clustered in a distinct but monophyletic clade with M. kontumensis from south-central Vietnam (Fig. 4). However, it has a genetic distance, based on COI, of 11.5% from M. kontumensis. This estimate evolutionary divergence is relatively large for intraspecific variation within Murina species, but it is typical between morphologically similar sister species, e.g. M. chrysochaetes vs M. harpioloides (11.9%; Eger & Lim, 2011). It is interesting to note that based on COI sequences the specimens of M. cf. eleryi from the same locality as the new species, although clustered closely with other specimens of M. eleryi, form a distinct clade and are more similar with M. balaensis from peninsular Thailand (K2P 9.3%) than with specimens from elsewhere in Indo-China (10.5%) (Fig. 4). It is very likely that M. eleryi has a complex biogeographic pattern, as mentioned in Son et al. (2015). Further study may demonstrate that the population of ‘ M. cf. eleryi’ in the Hkakaborazi Landscape has a distinct genetic structure, reflecting its demographic history, e.g. paleo-refuges, and that it is another undescribed cryptic species (P. Soisook, unpublished data). Comparison with similar species. Murina hkakaboraziensis sp. nov. differs from Murina species in the cyclotis- type in having relatively small P2. Among species with a suilla- type morphology, it can be separated from most species by a combination of its relatively small size (FA 29.6 mm; GTL 13.82 mm – Table 1 and 2), and pelage colour, having grey hairs on the belly and orange-brown hairs on the back with long golden guard hairs on the head and back. It is relatively similar in body size to M. harpioloides and M. chrysochaetes; it also shares the character of having golden hairs. However, it differs genetically and in other details of the pelage colouration and craniodental morphology. Murina harpioloides has a more orange colouration and the sagittal and lambdoid crests, although very weak, are present (Kruskop & Eger 2008; Son et al. 2015); they are absent in M. hkakaboraziensis sp. nov. In M. chrysochaetes, the ventral pelage has golden hairs as on the dorsal side (Eger & Lim 2011), but in M. hkakaboraziensis sp. nov., the golden hairs are only apparent on the dorsal side (Fig. 1). The slope of the forehead of M. chrysochaetes is abrupt and the braincase is high and dome-shaped (Eger & Lim 2011), but the forehead profile of the new species gradually slopes up to the braincase, which although relatively high, is not dome-shaped (Fig. 2a). The pelage appearance of the new species is also somewhat similar to Harpiola isodon and M. lorelieae, but both of them have a larger body size and differ in the shape of the cranium (Kuo et al. 2006; Eger & Lim 2011; Son et al. 2015). Murina hkakaboraziensis sp. nov. is most similar to the Vietnamese M. kontumensis, which was very recently described from south-central highland of Vietnam (Son et al. 2015). However, M. hkakaboraziensis sp. nov. has a more plentiful and well-defined longer bright golden guard hair (Fig. 1a). Besides the bright golden guard hairs, the colouration of the pelage on the dorsal and ventral sides are also different. In M. kontumensis, the general impression of the hairs are brownish-grey on the dorsal side and light brown on the ventral side, whereas M. hkakaboraziensis sp. nov., is orange-brown on the dorsal side (Fig. 1b) and whitish-grey on the ventral side (Fig. 1c). There is no contrasting whitish collar around the neck of M. hkakaboraziensis sp. nov. (Fig. 1a), but this whitish collar is very obvious in M. kontumensis (Son et al. 2015, figure 3A). The wing membrane of the new species is attached about 2 mm above the base of the claw of the outer toe, whereas it attached to the base of the claw in M. kontumensis (Son et al. 2015, figure 3H). In the cranium, the lateral profile of the interorbital region of M. kontumensis rises abruptly to the forehead, but rises smoothly and gradually in M. hkakaboraziensis sp. nov. (Fig. 2). The sagittal crest of M. kontumensis is well-defined but it is absent in the new species (Fig. 2). In the dentition, the secondary cusp of the I3 of M. hkakaboraziensis sp. nov. is very well developed and is about half the height of the main cusp, but in M. kontumensis this cusp is very poorly developed and is less than one-third the height of the main cusp (Fig. 2b). The posterior upper premolar (P4) of M. hkakaboraziensis is somewhat square (Fig. 2a) and larger in crown area than that of M. kontumensis, which is rectangular, broader than long (Fig. 2b). Subsequently, the upper canine (C1) of M. hkakaboraziensis sp. nov. appears to be relatively smaller in crown area than in M. kontumensis when comparing to the P4. Ecology and distribution. The new species, M. hkakaboraziensis sp. nov., was collected in a mist net set at the edge of a lowland semi-evergreen forest at the transition zone to an open space grassland, which undergoes an annual burn (Fig. 5). The new species was the only bat captured in the mist net. However, on the same night, four other insectivorous bats, Rhinolophus affinis, R. pusillus, Aselliscus stoliczkanus and Hipposideros pomona were captured in nearby mist nets and harp traps. Four other vespertilionids, M. cyclotis, M. feae, M. cf. eleryi , Kerivoula hardwickii, and K. furva were also captured in the same area on other nights. Currently, the new species is only known from the holotype collected from the type locality in the Hkakabo Razi Landscape, Kachin, northern Myanmar., Published as part of Renner, Swen C., 2017, A new species of Murina (Chiroptera: Vespertilionidae) from sub-Himalayan forests of northern Myanmar, pp. 159-172 in Zootaxa 4320 (1) on pages 161-166, DOI: 10.11646/zootaxa.4320.1.9, http://zenodo.org/record/891627, {"references":["Eger, J. L. & Lim, B. K. (2011) Three new species of Murina from Southern China (Chiroptera: Vespertilionidae). Acta Chiropterologica, 13, 227 - 243. https: // doi. org / 10.3161 / 150811011 X 624730","Son, N. T., Csorba, G., Tu, V. T., Thong, V. D., Wu, Y., Harada, M., Oshida, T., Endo, H. & Motokawa, M. (2015) A new species of the genus Murina (Chiroptera: Vespertilionidae) from the Central Highlands of Vietnam with a review of the subfamily Murininae in Vietnam. Acta Chiropterologica, 17, 201 - 23. https: // doi. org / 10.3161 / 15081109 ACC 2015.17.2.001","Kruskop, S. V. & Eger, J. L. (2008) A new species of tube-nosed bat Murina (Vespertilionidae, Chiroptera) from Vietnam. Acta Chiropterologica, 10, 213 - 220. https: // doi. org / 10.3161 / 150811008 X 414809","Kuo, H. - C., Fang, Y. - P., Csorba, G. & Lee, L. - L. (2006) The definition of Harpiola (Vespertilionidae: Murininae) and the description of a new species from Taiwan. Acta Chiropterologica, 8, 11 - 19."]}

Bayesian clustering methods are typically used to identify barriers to gene flow, but they are prone to deduce artificial subdivisions in a study population characterized by an isolation-by-distance pattern (IbD). Here we analysed the landscape genetic structure of a population of wild boars (Sus scrofa) from south-western Germany. Two clustering methods inferred the presence of the same genetic discontinuity. However, the population in question was characterized by a strong IbD pattern. While landscape-resistance modelling failed to identify landscape features that influenced wild boar movement, partial Mantel tests and multiple regression of distance matrices (MRDMs) suggested that the empirically inferred clusters were separated by a genuine barrier. When simulating random lines bisecting the study area, 60% of the unique barriers represented, according to partial Mantel tests and MRDMs, significant obstacles to gene flow. By contrast, the random-lines simulation showed that the boundaries of the inferred empirical clusters corresponded to the most important genetic discontinuity in the study area. Given the degree of habitat fragmentation separating the two empirical partitions, it is likely that the clustering programs correctly identified a barrier to gene flow. The differing results between the work published here and other studies suggest that it will be very difficult to draw general conclusions about habitat permeability in wild boar from individual studies.

The northern Myanmar hill jungle and neighboring highlands found along the southeastern slopes of the Himalayas (Hkakabo Razi region, northern Kachin State, northernmost Myanmar) is almost untouched by humans, and can be considered as primary forest in the literal sense. Until the efforts reported on in this monograph, no ornithologist had the opportunity to visit this remote area since the late 1940s. Beginning in 1997, we made several visits to this extraordinary, species-rich area, and compiled an inventory of the regional avifauna while also studying the ecology, behavior, systematics, taxonomy, biogeography, and origin of the region’s birds. We report on these aspects for the 441 species found in the area and add data on species new to the area and those for which there are only a few representatives in the world’s specimen collections. In addition, we provide information on taxa that are likely endemic (Jabouilleia naungmungensis), suggest splits or revisions of subspecies and species, and describe two new subspecies from the area, Alcippe cinereiceps hkakaboraziensis ssp. nov. and Malacocincla abbotti kachinensis ssp. nov., based on plumage and morphometric differentiation. Last but not least, we analyze the species affinities (i.e. the biogeographic origin of the species occurring in Hkakabo Razi and surrounding areas). We found a lack of data in general for bird species in Southeast Asia, India and Tibet/China, but especially in the northern hill mountains of Kachin State around Hkakabo Razi National Park. Efforts by conservation organizations and ourselves has improved the situation, but much more work, particularly on ecology and global change effects, is warranted. We have added considerable information on the distribution, systematics, ecology, and biogeography of a number of species. Although much work remains to be done, our research provides significant new insights into the biology and biogeography of the region’s birds, and in particular, on delineation of major zoogeographic regions. We conclude that Ernst Mayr was correct in essence when he stated that the eastern border between the Oriental and Southeast Asian regions is roughly equivalent to the political boundary between Myanmar (Burma) and China. However, our work provides new understanding of the reasons, and challenges some of the fundamental assumptions on which the ‘zoogeographic region’ concept is based. In particular, such delineations often fail to take into account the geological and ecological history of a region. By evaluating in detail the subspecific relationships of the Hkakabo Razi rainforest avifauna, we were able to determine that the avifauna most similar is found in the temperate rainforests of northeastern India. We conclude that the likely reason for this similarity is that these two avifaunas shared the same refugium during the maximum of the last glacial period (18,000 years ago), and that other south and southeast Asian rainforest species shared different refugia during this time period, and probably during previous glacial events as well., Other parts of this work have been funded through six SYNTEHSYS grants (visits to BMNH, NMN, NMW, ZMUC, NRM, and Museo de la Historia Natural de Madrid: ES-TAF-2501; AT-TAF-2481; GB-TAF-108, SE-TAF-1312; NL-TAF-4369, AT-TAF-7), under the European Community’s 7th Framework Program (www.synthesys.info).

Many studies have examined whether communities are structured by random or deterministic processes, and both are likely to play a role, but relatively few studies have attempted to quantify the degree of randomness in species composition. We quantified, for the first time, the degree of randomness in forest bird communities based on an analysis of spatial autocorrelation in three regions of Germany. The compositional dissimilarity between pairs of forest patches was regressed against the distance between them. We then calculated the y-intercept of the curve, i.e. the 'nugget', which represents the compositional dissimilarity at zero spatial distance. We therefore assume, following similar work on plant communities, that this represents the degree of randomness in species composition. We then analysed how the degree of randomness in community composition varied over time and with forest management intensity, which we expected to reduce the importance of random processes by increasing the strength of environmental drivers. We found that a high portion of the bird community composition could be explained by chance (overall mean of 0.63), implying that most of the variation in local bird community composition is driven by stochastic processes. Forest management intensity did not consistently affect the mean degree of randomness in community composition, perhaps because the bird communities were relatively insensitive to management intensity. We found a high temporal variation in the degree of randomness, which may indicate temporal variation in assembly processes and in the importance of key environmental drivers. We conclude that the degree of randomness in community composition should be considered in bird community studies, and the high values we find may indicate that bird community composition is relatively hard to predict at the regional scale.

Am 08. Januar 2007 gegen 11:00 Uhr morgens konnte ich in der Wuppertaler Innenstadt (Stadtteil Elberfeld: 51° 16‘ 26“ N, 007° 08‘ 41“ O, ca. 150 m ü NN) ein Amselweibchen (Turdus merula) beim Füttern von mindestens zwei sperrenden Nestlingen beobachten. Das Nest war auf der Leuchtreklame eines größeren Modehauses unter einem Glasdach platziert. I observed a breeding attempt of an European Blackbird Turdus merula in the city of Wuppertal (North Rhine Westphalia, western Germany: 51° 16’ 26” N, 007° 08’ 41” E, 150 m a.s.l.) on 08 January 2007. The female Blackbird was feeding at least two nestlings. The nest was placed above a luminous advertising above an entrance to a larger fashion store. The nest was covered by glass roofing above the entrance to the store.