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Remaining Cryptic During Motion — Behavioral Synchrony in the Proboscis Bat (Rhynchonycteris Naso)

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Remaining Cryptic During Motion — Behavioral Synchrony in the Proboscis Bat (Rhynchonycteris Naso) Acta Chiropterologica, 11(1): 208–211, 2009 PL ISSN 1508-1109 © Museum and Institute of Zoology PAS SHORT NOTES doi: 10.3161/150811009X465839 Remaining cryptic during motion — behavioral synchrony in the proboscis bat (Rhynchonycteris naso) MIRJAM KNÖRNSCHILD1, 4, CHRISTINA HARVIEW2, RACHEL MOSELEY3, and OTTO VON HELVERSEN1 1Department of Zoology, University Erlangen-Nürnberg, Staudtstrasse 5, D-91058 Erlangen, Germany 2Department of Biology, Bryn Mawr College, 101 N. Merion Avenue, Bryn Mawr, PA 19010, USA 3Department of Biology, Bucknell University, 701 Moore Avenue, Lewisburg, PA 17837, USA 4Corresponding author: E-mail: [email protected] Key words: Rhynchonycteris naso, cryptic roosting, behavioral adaptation, synchronous rocking, grooming, urinating INTRODUCTION individuals of both sexes (Bradbury and Vehren - camp, 1976). Individuals of small groups normally Most bat species inhabit roosting sites in which roost in a vertical line, whereas larger groups tend to they are well concealed during the day, such as form ovals (Bradbury and Emmons, 1974). Indepen - caves or tree holes, whereas others roost in exposed dent of group size, a vertical line is always formed locations on tree trunks or branches (Kunz and Lum - when individuals roost on structures with a small sden, 2003). For the latter, it is essential to appear diameter, such as thin trunks, branches and vines cryptic in order to avoid predation or disturbance at (authors’ personal observation). Despite its exposed the day-roost. Several traits have evolved to enhance day-roosts, R. naso is physically very cryptic due to the bats’ cryptic appearance, including cryptic col- the woolly and mottled pelage with two wavy pale oration (e.g., the greenish wings of the foliage-roost- lines on the back and tufts of pale hair on the fore- ing bat Paranyctimene raptor — Bona ccor so, 1998) arms (Dalquest, 1957; Bradbury and Vehrencamp, or disruptive markings (e.g., the contrasting meta - 1976). This protective camouflage allows the roost- car pals and phalanges of the tent-roosting flying ing bats to resemble a patch of lichen or moss (Hill fox Cynopterus brachyotis — Kunz and Lumsden, and Smith, 1984). Bradbury and Emmons (1974) 2003). Behavioral habits may also aid in camouflag- first reported a ‘synchrony in grooming periods by ing roosting bats. One remarkable example is the ‘leaf wrapping behavior’ exhibited by the Australian group members’ and ‘frequent periods of gentle flute-nosed bat Murina florium, which conceals rocking’, a stereotypic behavior in which the bats these foliage-roosting bats effectively from visually sway rapidly from side to side while both feet and oriented predators (Schulz, 1999). In general, there wrists remain attached to the surface of the roost. are many more known examples of morphological This peculiar behavior occurs frequently during the adaptations for cryptic roosting than there are of day and is often exhibited synchronously by many behavioral adaptations (Kunz, 1982; Kunz and or all group members, but it is not triggered by dis- Lum sden, 2003).Remaining completely motionless turbance. When startled or approached, the bats do during the day is a behavioral habit that obvious- not rock, but instead remain motionless before they ly assists in camouflage, but it conflicts with the fly off together. Therefore, the functional signifi- bats’ need for comfort behavior, such as grooming cance of rocking has remained unclear (Bradbury or stretching. and Emmons, 1974), but a somewhat related behav- We studied the cryptic roosting habits of the pro- ior in arachnids, termed whirling or bobbing, is pro- boscis bat, Rhynchonycteris naso. This neo-tropical posed to provide a camouflage defence against pred- member of the emballonurid family roosts in very ators (Jackson, 1990; Heuts et al., 2001; Grether and exposed areas on tree trunks, branches, vines or Donaldson, 2007). Group members of R. naso not man-made structures which are in close proximity only synchronize rocking and grooming but also uri- to running water (Bradbury and Emmons, 1974). nating (authors’ personal observation), but the bene- Group size is intermediate, and ranges from 5 to 50 fit, if any, of this behavioral synchrony is unknown. Short Notes 209 We investigated whether rocking could play tests were conducted using SPSS version 11.5 (SPSS Inc., a role in the bats’ cryptic roosting habits. We firstly Chicago, IL, USA). hypothesized that bats engage in rocking behavior to resemble patches of vegetation swaying in the wind RESULTS and, hence, that rocking is triggered by gusts of wind. We tested this hypothesis by relating the oc- Rocking occurred significantly more often dur- currence of rocking to simultaneous wind velocity at ing a gust of wind than during a calm (repeated the roosting sites. We secondly hypothesized that measures ANOVA: F1, 4 = 238.61, P < 0.001; mean synchronous rocking by many individuals is used to oc currence of rocking during gusts and calms for actively disguise other behaviors, such as grooming every observation session; five groups with 10 re- or urinating, and tested this hypothesis by conduct- peated measures each). Nonetheless, the relation be- ing behavioral observations during rocking periods. tween the occurrence of rocking and wind velocity was not mandatory: not every gust of wind was ac- MATERIALS AND METHODS companied by rocking (143, i.e. 34.6% of the 413 gusts measured were not followed by rocking; wind Observations and measurements were carried out at the velocity: 0.1–0.5 m/s) and, likewise, single bats Biological Station La Selva in Costa Rica (10°25’N, 84°00’W) sometimes rocked briefly despite a complete calm in two consecutive years (July 2007, June 2008). The five social groups of R. naso that were used for this study occupied day- (53, i.e. 11.4% of the 466 rocking occurrences ob- roosts under the roofs of both abandoned and inhabited build- served). We found a significant positive relation be- ings throughout the station property. These ‘artificial’ day- tween the wind velocity and the percentage of rock- roosts were comparable to ‘natural’ day-roosts (e.g., on tree ing bats per group (GLMM with wind velocity as in- trunks) with regard to their exposure to general weather condi- dependent and % of rocking bats as dependent vari- tions like heat and wind. Group size ranged between 8 and 35 bats. Bats of all groups were habituated to the presence of hu- able: F1, 19 = 521.00, P < 0.001). An increase in man observers, which made it possible to gather data without wind velocity by 0.2 m/s caused an approximate causing noticeable disturbance. 25% increase in the amount of rocking bats per Each group was observed ten times for a minimum of group (fixed effect: wind velocity; estimate = 0.246, 30 minutes per observation session (total observation time: T = 22.83, P < 0.001; see Fig. 1A). 37.4 hours). During each observation session, we recorded the 19 occurrence of rocking and the associated behaviors grooming We found a significant positive relation between and urinating. We estimated the total number of bats rocking by the percentage of rocking bats per group and the oc- assigning one of five categories (0, 25, 50, 75, and 100% of currence of both grooming and urinating (GLMM the group). This estimation was conducted at the end of each with % of rocking bats as independent variable and rocking bout because usually one or few individuals started (a) occurrence of grooming as dependent variable: rocking and then others progressively joined in. When a single bat rocked for only a few seconds and ceased without others F1, 23 = 20.37, P < 0.001; and (b) occurrence of joining in, we assigned the 0% category because we interpreted urina ting as dependent variable: F1, 23 = 23.93, it as an unsuccessful attempt of one individual to engage other P < 0.001). Synchronous grooming increased con- group members in rocking behaviour. We also noted whether stantly as the percentage of rocking bats per group rocking was accompanied by either grooming (i.e., licking and/or scratching of pelage and wing membranes) or urinating increased, whereas synchronous urinating occurred (i.e., lifting the body away from the surface with both forearms only when at least 75% of all group members exhib- and feet while excreting clearly visible droplets of urine) or ited rocking behavior (Fig. 1B). In the majority of both. In addition, wind velocity measurements were conducted ca ses, rocking was accompanied by either groom- every two minutes and whenever rocking occurred. We used ing or urinating (rocking with associate behav- a Kestrel® 1000 Pocket Wind Meter (Nielsen-Kellerman Co., Boothwyn, PA 19061, USA; range: 0.1–60.0 m/s; resolution: iors = 56%; rocking without associate behav- 0.1 m/s) that was placed in close vicinity to the roosting bats iors = 38%; grooming without rocking = 6%). This (maximum distance of two meters). suggests that synchronous rocking is used to conceal Since multiple rocking periods could occur during an obser- other behaviors which, undisguised, might under- vation session, the data was averaged per category (0, 25, 50, mine the cryptic roosting habits of R. naso. 75, and 100% of the group) for each session. The occurrence of grooming and urinating was scored with values of 0 or 1 for each rocking bout, added for all the rocking bouts within an ob- DISCUSSION servation session and then averaged over all observation ses- sions per group (always separately for each rocking category). Our study shows that proboscis bats are more We used General Linear Mixed Models (GLMM) to test for re- lations between wind velocity, rocking, and associate behaviors. likely to rock during gusts of wind than during Multiple measurements per group were controlled for by adding calms, and that the proportion of rocking group group identity as a random factor in the GLMM.
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