MARINE MAMMAL SCIENCE, 15(4):1065-1097 (October 1999) © 1999 by the Society for Marine Mammalogy DIURNAL BEHAVIOR OF FREE-RANGING | BOTTLENOSE DOLPHINS IN THE KVARNERIC (NORTHERN ADRIATIC SEA)! Giovanni Bearzi tena Pour ‘Tethys Research Institute, viale G.B. Gadio 2, I-20121 Milano, Italy E-mail: bearzi@tin.ic Gruseppz NoTARBARTOLO DI SCIARA ICRAM, via Casalotti 300, 1-00166 Roma, Italy Apstaact ‘The diurnal behavior of a bottlenose dolphin (Tersiops truncatus) community ‘was observed from small inflatable craft between 1987 and 1994. Following a preliminary ad libitum study 11,839 3-min behavioral samples were recorded in 1991-1994, The behavioral budget showed a predominance (about 80%) of activities characterized by long (>30 sec) dives, considered to be largely related to prey search or feeding. Obvious foraging near the surface was ob- served rarely. The frequent following of trawlers (accounting for 4.6% of the behavioral budget) was indicative of the presence of alternative strategies for finding food. Yearly and seasonal behavioral variation—particularly in feed- ing-telated and travel behaviors—was consistent with the hypothesis of be- havioral flexibility as a response to environmental changes and fluctuating prey kind and availability. Yearly shifts in social behavior appeared co be partly influenced by breeding cycles. Groups engaged in feeding-related ac- tivities were significantly smaller than traveling or socializing groups, and dramatic interannual group-size shifts seemed to be largely affected by en- vironmental variables, rather than being entirely determined by behavioral activity changes. The remarkable behavioral flexibility of this bottlenose dol- phin community may contribute to its survival in che shifting environmental Conditions of the northern Adriatic Sea. However, the high proportion of time consistently devoted to feeding-related activities, as compared to other areas, suggests that food resources in the Kvarnerié were not only highly variable but also depleted. Key words: bottlenose dolphin, Tursiops truncatus, Mediterranean Sea, Adriatic Sea, behavior, group size. "We dedicate chis work to Ken Notts, who first gave che statute of solid science to the act of observing wild dolphins: in deep appreciation for entiching us with the inspiration of his teaching, with the example of his imaginative work, and with the encouragement of his ever ftiendly and liberal advice.1066 MARINE MAMMAL SCIENCE, VOL. 15, NO. 4, 1999 Despite a number of studies focusing on the social ecology of coastal dolphin communities and a proliferation of descriptive behavioral accounts, the behav- ior of free-ranging bottlenose dolphins has been the subject of few quantitative investigations, these mostly conducted in shallow inshore waters (Shane ¢f al. 1986; Shane 1990z, 6; Bassos 1993; Hanson and Defran 1993). This is hardly astonishing, as the quantitative description of the behavior of often elusive, deep-diving dolphins which are visible at the surface only for a small per- centage of the time can be a difficult task. However, data obtained from a small boat, causing minimal disturbance to the animals, are especially valuable when systematically collected through standard procedures and associated with results from alternative approaches (e.g., acoustic investigations). In this long- term study conducted near che Croatian islands of Loinj and Cres, where the conditions did not allow for either underwater observations or safe captures (@g., for tagging or for radio-tracking), a non-invasive observational approach based on behavioral sampling was adopted, in association with systematic photoidentification (Bearzi et al. 1997), focal-individual respiration sampling (orcuna 1996), and acoustic data collection (Ochen 1996). As the research continues, combined data analysis are providing insight into the social ecology and behavior of this bottlenose dolphin community, which lives in a sea that has faced environmental degradation, dramatic eutrophication phenomena, and shifts of prey kind and availability during recent decades (Bombace 1990, Chiaudani and Premazzi 1995, Marasovié ef al. 1995, Cingolani et al. 1996, Conti 1996, UNEP 1996, Della Croce e al, 1997, Soli¢ et al. 1997). The possibility of defining and promoting effective conservation measures, to pro- tect the only cetacean species now consistently observed throughout the north- etn Adriatic Sea (Notarbartolo di Sciara and Bearzi 1992, Bearzi and Notar- bartolo di Sciara 1995, Gomertié et al. 1998), rests on understanding the survival problems faced by the dolphins. ‘Marraiats AND MeTHops The study area, roughly 800 km?, is situated in the southern portion of the Kvarnerié (Fig. 1). The sheltered and clear coastal waters east of the islands of Lofinj and Cres, where most of the data were collected, encompass a variety of marine habitats, including rocky coastline and bottom, submerged reefs, seagrass flats, and a mud sea bed. The typical depth is about 70 m and the maximum depth 96 m. The water temperature undergoes wide yearly varia- tion, with surface values ranging between 10° and 30°C. Water clarity, mea- sured by Secchi disk, reached 20 m. Seasons were defined as spring: March— May, summer: June-August, fall: September-November, and winter: Decem- ber-February Ghane 19902, Bearzi et al. 1997). Observations were performed from September 1987 to September 1994, with a coral of 38 mo spent in the field (Fig. 2). Ad Jibitum surveys were conducted from inflatable boats with fiberglass keels, equipped with 25-45 HP outboard engines. An even distribution of surveys was attempted in order to cover all parts of the study area every month, although the geographicBEARZI £7 AL.: DIURNAL BEHAVIOR OF BOTTLENOSE DOLPHINS 1067 [Fase | \ a By Py Figure 1. Study area, distribution of the effort could vary due to weather conditions and dolphin sighting frequency (Bearzi ef a/. 1997). A total of 388 d were spent at sea in 1987-1994, totalling 752 h spent observing and photographing 787 dolphin groups encountered during 311 uninterrupted observations lasting on average 143 min (range 15-615 min). In order to evaluate the occurrence of dolphins in the proximity of bottom trawlers, 202 boats engaged in trawling (i.e. dragging the net at an average speed of 2~4 kn) were opportunistically ap- proached when encountered during navigation in search for dolphins in 1990— 1994, The presence of dolphins was then verified by stopping the boat near the wake of the fishing boat, at about 200 m from its stern, and carefully observing the sea surface for a minimum of 5 min. In 1987-1990 behavioral data were collected ad libitum, thus providing a framework of information (160 h of observation) to design the behavioral sampling procedures that were systematically adopted from 1991. Data col- lected in 1991-1994 (326 d at sea, 592 h of observation) represented the primary dataset for behavioral analyses. Across this sampling period, behavioral data were routinely recorded at the end of 11,839 standard 3-min intervals (Alemann 1974), focusing on 688 different dolphin groups, with a mean of