Anti-predator behavior in : Reactions to Raptor Calls

Robin Elahi

Department of Biology, Northeastern University

______ABSTRACT

Birds display a number of defense mechanisms against their predators, including camouflage, mobbing, fleeing, and aggregational behavior. I studied the responses of 32 to songbird and raptor calls to see if birds displayed anti-predator behavior in response to raptor calls in comparison with songbird calls. One focus of my study was avian defensive strategies, namely flocking and concealment, so observations took into consideration different conditions: whether the bird was solitary or in a group, and whether it was exposed or hidden in vegetation. A previous study showed that crows could distinguish between raptor calls of their predators and non-predators, so I examined if birds specifically recognized resident raptors, or if they had a general response to resident and non-resident raptor calls. I also attempted to correlate bird size with reaction, specifically to see if smaller birds flew away more often than larger birds due to a greater perceived threat. For each bird or group of birds. I played a songbird call, followed by a raptor call, and timed the raptor call to see how long the bird stayed till it flew away. Birds flew away 27% of the time in response to raptor calls, and only 3% of the time to songbird calls (X² = 16.11, df = 1, n = 26). Birds did not distinguish among resident and non-resident raptor calls, and had nearly equal flight frequencies (X² = 0.48, df =3, n=40). Trends showed that solitary and exposed birds tended to fly more often than birds in groups and hidden birds. Bird size was not a reliable indicator of reactions, although a trend demonstrate that average bird size was larger for flight reactions, than for no reactions or look reactions (One-way ANOVA, p = 0.1314). These results suggest that birds use the anti- predator strategies of flocking and concealment, however, further research should be done on raptor call recognition on a single or species of birds to determine if raptor calls recognition is specific on a lower taxonomic level as previous studies have suggested.

RESUMEN

Los pájaros exhiben una variedad de mecanismos defensivos contra sus predadores, incluyendo camuflaje, ataque, huir o hacer grupos. Estudie las reacciones de 32 especies de pájaros al sonar cantos de aves canoras y aves de rapiña, para ver si los pájaros exhibieron un comportamiento anti-predador. El enfoque de mi estudio fueron las estrategias de las aves, específicamente en grupos y escondidos, por eso en las observaciones se tomaron en consideración si los pájaros estuvieron en grupo o solitarios o si estuvieron escondidos en la vegetación o no. Un estudio anterior demostró que los cuervos distinguieron entre las llamadas de rapaces, por eso examine si los pájaros específicamente distinguieron entre llamadas de aves de rapiña residente y no-residente, o si tuvieron una respuesta general a todas las llamadas de ave de rapiña. También trate de tomar en cuenta el tamaño del pájaro con la reacción, específicamente si los pájaros pequeños volaron más que los pájaros grandes. Para cada pájaro o grupo de pájaros, soné una llamada de ave canora e inmediatamente después una llamada de ave de rapiña y conté el tiempo que le tomo al pájaro volar. Los pájaros volaron en 27% del tiempo en respuesta a las llamadas de rapiña y solamente 3% del tiempo a las llamadas de aves canoras (X² = 16.11, df = 1, n = 26). Los pájaros no distinguieron entre las aves de rapiñas residentes y no-residentes y tuvieron casi la misma frecuencia de vuelo (X² = 0.48, df = 3, n = 40. Las tendencias mostraron que pájaros solitarios y expuestos volaron más que las aves en grupo o escondidas. El tamaño del pájaro no fue un indicador para las reacciones pero una tendencia mostro que el tamaño fue un factor para cuando volaron, que para cuando no lo hicieron o para cuando solo miraron (One-way ANOVA, p= 0.1314). Estos resultados sugieren que los pájaros usan estrategias anti-predador, pero se debe estudiar más el reconocimiento de una especie de pájaro de cómo distingue las llamadas de aves de rapiña para determinar si el reconocimiento es específico en especies como mostraron los estudios anteriores.

INTRODUCTION

Predators confer Strong selective pressure on their prey, resulting in varied defense mechanisms. have developed two basic types of strategies to prevent predation – anti-detection and anti-capture (Alcock 1984). Anti-detection strategies rely on remaining invisible to the predator, through cryptic coloration and background matching. This is seen in many different taxa, such as lizards, insects and birds. Anti-capture strategies are behaviors displayed after the prey has been spotted by their predator, such as flash colors, misdirecting an attack, chemical repellants, Batesian mimicry, fleeing and fighting back (Alcock 1984). Birds avoid predation by raptors using both anti-detection and anti-capture strategies. Many birds, such as Amazonia parrots, blend in with their environment and are very difficult to see, which serves as an anti-detection strategy. Once a raptor has spotted its prey, anti-capture strategies include diving into thick bushes or water, and simply fleeing and dodging. These actions are often accompanied by alarm calls (Perrins 1979). Birds have also evolved anti-capture behaviors such as mobbing and aggregational behavior. For example, European kestrels mobbed by foraging birds flew significantly farther from foraging areas, and thus supports the assumption that mobbing decreases predatory efficiency, mainly by driving the predator from the vicinity (Pettifor 1990). Aggregational behavior has been shown to be an effective defense against predators in numerous studies on leks, foraging flocks, and escape tactics (Buchanan et al. 1988, Caldwell 1986, Glodman 1980, Trail 1998). Flocks provide increased vigilance because of the greater number of eyes to spot predators. As a result, individuals can increase the time they spend foraging since they spend less time watching for danger. Lima (1998) has shown that solitary birds that are not overtly vigilant (i.e. birds that are feeding) are more vulnerable to predator attack. Additionally, the selfish-herd hypothesis states that an individual’s chances of being eaten decrease when they join a group, simply because of probability (Terborgh 1989). The dilution effect explains that groups will likely satiate their predators, and again, the individual’s chances of survival increase due to probability, since only a few prey will be taken (Alcock 1984). Since aggregational behavior and concealment are key defense mechanisms for birds, I tried to measure the effects of these variables on the reaction of birds in response to songbird and raptor calls. The reactions to songbird calls provide a control to compare the reactions of raptor calls. Based on avian anti-predator strategies, solitary birds should fly away more often and more readily than birds in groups. Likewise, exposed birds should fly away more often than hidden birds. I am also interested in whether birds respond specifically to the calls of resident raptors, or whether they have a general raptor call response. Western American Crows (Corvus brachyrhynchos hesperis) not only distinguish between raptor and non-raptor calls, they can also distinguish the calls of Red-Shouldered Hawks ( lineatus), a major predator, from the Madagascar Harrier Hawk (Gymnogenys radiatus) that has a spectrally similar call (Hauser and Caffrey 1994). Therefore, it appears birds will distinguish between songbirds and raptors, and that at least some birds will recognize the calls of raptors that are common in the area, and as a result, fly more often and more readily from those calls. The Great Blawk-Hawk ( anthracinus) and the Collared Forest-Falcon (Micrastur semitorquatus) were chosen as representative of the San Luis area, while the Semiplumbeous Hawk ( seminplumbea) and the Slaty-Backed Forest-Falcon (Micrastur mirandollei) frequent other areas, mainly the Caribbean slope (Mauricio Ramirez, Manuel Leitón, pers. comm.; Fogden 1993, Stiles and Skutch 1989). As a last inquiry, I wished to examine the relationship between bird size and reaction. Only larger birds were found to mob kestrels because it seemed unlikely that the kestrels posed a great threat to them (Pettifor 1990). I hypothesize that smaller birds will fly more often than larger birds, because of the greater perceived threat.

MATERIALS AND METHODS

Calls were played to 87 birds of 32 different species in San Luis Arriba and Invu, Costa Rica, located on the Pacific slope of the Tilarán mountain range at 1100m elevation between October 25 and November 14, 2000. The Holdridge life zone is premontane wet forest (Holdridge 1967, Haber et al. 2000). Data were collected from 5:30 – 8:30 AM and 3:00 – 5:00 PM, along roads and trails in pasture, secondary growth forest and gardens. I walked trails, stopping to make observations at certain points for durations of five to twenty minutes depending on bird activity. Once a bird was perched and identified using binoculars, I played a call sequence once, at medium to high volume depending on distance from the observed bird (i.e. volume increased with increased distance) with a portable cassette player, pointed straight up in the air. The distance ranged from two meters to 25m. There were four different call sequences, each consisting of a songbird call, immediately followed by a raptor call, as follows (time in seconds is noted in parentheses): Sequence 1: Tropical Pewee (39s), Great Blawk-Hawk (39s) Sequence 2: Buff-Throated Wood-Creeper (44s), Semiplumbeous Hawk (35s) Sequence 3: Striped-Breasted Wren (45s), Collared Forest-Falcon (54s) Sequence 4: Yellow-Faced Grassquit (29s), Slaty-Backed Forest-Falcon (33s) The specificity of bird responses to native raptors was tested by sequences one and three, and to non-native raptors by two and four, as explained above. At the start of the raptor call I started timing to see how long the bird stayed. I categorized three reactions to the bird calls – “None”, “Look”, and “Fly”. “None” indicates the bird did nothing by the time call ended. “Look” meant the bird stopped his current activity (e.g. singing, pecking) and looked around, and “Fly” meant the bird flew away. Each call sequence was played only once. If the bird was by itself, it was “Solitary”, and if it was with at least one other bird (only one group of birds I observed was made up of more than one species) that was noted as a “Group”. Groups that displayed a “fly” reaction had at least half of their birds fly away. Whether the bird was “Exposed” or “Hidden” was also noted – “Exposed” meant that the bird was in clear view; if the bird was at least partly surrounded by vegetation it was defined as “Hidden”. The birds I chose for songbird calls are in the order Passeriformes. This order is characterized by the morphology of the syrinx (the organ of sound production) (Janzen 1983). The four chosen songbirds range from 10 to 22 cm in length, and eat a variety of seeds, fruits and insects. The raptors whose calls I used range from 38 to 66cm; they eat birds and eggs (Stiles and Skutch 1989). The raptors are in the order Falconiformes, which includes hawks, and falcons. They are diurnal hunters who rely mainly on sight; they have no sense of smell (Perrins 1979). These birds of prey are vocal when demonstrating territoriality, and when performing courtship rituals, as most other birds (Perrins 1979). Skutch and Stiles (1989) aided in identification, and reported bird size in length (cm). Chi square tests were used to determine the significance of bird reactions to the different calls (songbird vs. raptor, raptor vs. raptor), and under the four different conditions (solitary vs. group, exposed vs. hidden). One-way ANOVA tests were used to relate flight times with the different raptor calls and the different conditions. One-way ANOVA tests were also run between bird size and reaction.

RESULTS

Eighty-seven call sequences were played to birds; three of the birds flew after hearing the songbird calls, and thus 87 birds listened to songbird calls and 84 birds listened to raptor calls. There were significant differences between the songbird and raptor calls in all three reactions, “None”, “Look”, and “Fly” (X² = 30.78, df = 1, n = 99; X² = 26.57, df = 1, n = 46; X² = 16.11, df = 1, n = 26, respectively); (Figure 1). Differences in reaction were strikingly clear, for example, songbird reactions elicited no reaction 90% of the time, while raptor reactions elicited no reaction only 25% of the time. All four raptor calls were played to 21 birds each. The number of flight reactions were almost equal for all four calls (five for both the Great Blawk-Hawk and Collared Forest-Falcon, six for the Semiplumbeous Hawk, and seven for the Slaty-Backed Forest- Falcon). There was no significant differences in any of the reactions between the four raptor calls (X² = 1.29, df = 3, n=21, X² = 1.0, df = 3, n = 40, X² = 0.48, df = 3, n=40; “None”, “Look”, “Fly”, respectively); (Figure 2). For those birds that flew off, there was no significant difference in time (One-way ANOVA, p = .3019); (Figure 3). Raptor calls were played to 51 solitary birds, and 33 groups of birds. Flycatchers, such as Tropical Pewees (Contopus cinereus), Great Kiskadees (Pitangus sulphuratus) and the Myiozetetes flycatchers tended to be solitary, while Yellow-Throated Euphonias (Euphonia hirundinacea), Groove-Billed Anis (Crotophaga sulcirostris), and White- Fronted Parrots (Amazona albifrons) were always observed to be in groups. 35% of solitary birds flew, while only 15% of the groups flew (Figure 4) (X² = 2.98, df = 1, n = 23). For those that flew, that average time for solitary birds was 19.86 seconds, and for groups was 24.40 seconds (One-way ANOVA, p = 0.454). No reaction was observed 20% of the time for solitary birds, and 33% of the time for groups (X² = 1.51, df = 1, n =22). “Look” reactions were observed 45% and 52% of the time for solitary birds and groups, respectively (X² = 0.17, df =1, n=40). Raptor calls were played to 66 exposed and 18 hidden birds and groups (Figure 5). 32% and 11% of the subjects flew, respectively (X² = 2.20, df = 1, n = 23). The average flight time for exposed and hidden birds that flew was 21.69 and 12 seconds, respectively (One-way ANOVA, p = 0.271). No reaction was observed 21% of the time for exposed birds, and 39% of the time for hidden birds (X² = 1.77, df = 1, n = 21). “Look” reactions were observed 45% and 52% of the time for exposed and hidden birds, respectively (X² = 1.39, df = 1, n = 40). A significant X² value of 17.1 was calculated for a table between the solitary and group conditions versus the exposed and hidden conditions (Table 1). The exposed row contained 21 flight reactions, and the solitary column contained 18 flight reactions, while the hidden and group rows contained only two and five flight reactions respectively. Birds ranged from 10 cm (House wren) to 101 cm (Great Egret). When all the reactions to raptor calls were compared by bird size, there was no significant difference (One-way ANOVA, p = 0.846). To eliminate any bias from the different conditions (e.g. solitary vs. group), the test was run only for solitary and exposed birds, since that was largest sample size (n= 37); (One-way ANOVA, p = 0.131). The average bird size for fly reactions under these specific conditions was 21.47cm, while the average bird size for look and no reactions were 16.84 and 16.67, respectively. The difference lay between fly and look (One-way ANOVA, p = 0.064), and between fly and none (One-way ANOVA, p = 0.149) reactions.

DISCUSSION

Birds showed significant differences for all reactions with songbird calls and raptor calls (Figure 1). Ninety percent of the time, subjects showed no reaction to the songbird calls, compared with only 25% that showed no reaction to the raptor calls. The songbird calls thus served well as a control, demonstrating that the birds were not scared of me, and were not just reacting to sounds coming out of my tape player. Similarly, since birds only flew away 3% of the time to songbird calls, and 27% of the time to raptor calls, it suggests that birds did indeed perceive a difference between the two types of calls. These results support the previous study in which Western American Crows reacted to predator calls, but not to non-predators (Hauser and Caffrey 1994). The resident and non-resident raptor calls elicited nearly equal flight reactions (Figure 2). The times for flight were also not significantly different (Figure 3). Thus, it seems that birds have general raptor call recognition, instead of only recognizing specific raptors. This makes sense evolutionarily, because those birds who react to strange raptor calls would have higher fitness than those birds who only responded to familiar raptor calls. My results are different from Hauser’s and Caffrey’s (1994); they found that crows fly away only from their actual predator calls. The discrepancy here may be explained by the fact that I tested any species of bird I saw, while they studied one specific species, and they knew the actual predators from past research. The raptors whose calls I used are in the area, and they eat other birds and eggs in general, but it is unknown if they specifically search for prey of a certain genus or guild in the San Luis area. The birds that tended to be solitary and in groups as mentioned above confirmed the habits described in Skutch and Stiles (1989). The trend towards higher flight frequency and faster flight times among many species of solitary birds compared with birds in groups supports the hypothesis that solitary birds are more vulnerable to predation (Buchanan et al. 1988, Lima 1998). Since solitary birds are more vulnerable, and spend more time assessing their surroundings (Goldman 1980), it seems likely and more adaptive that they would flee more often. Being in groups is a safety measure due to increased vigilance, the selfish-herd hypothesis, and the dilution effect (Alcock 1984), and therefore the groups of birds did not find it necessary to flee. However, Trail (1987) found that larger mating groups, or leks, were actually more likely to get “spooked”, that is, to make head long flights, usually due to false alarms. The difference here may be due to the specific nature of leks. Since leks are groups of males dancing and singing, trying to attract females’ attention, the males would need to be very careful they weren’t attracting predators as well. In addition, his study was done in a tropical rain forest in Suriname, with a wide variety of predators, including raptors, felids and snakes. My hypothesis that exposed birds are more likely to fly than concealed birds was supported by the trend of 32% exposed birds flying due to raptor calls, compared with 11% of hidden birds flying. It is unusual that exposed birds had a higher flight time than hidden birds; however, only two hidden birds actually flew, so the average is not reliable. Also, hidden birds did nothing 39% of the time, compared with 21% for the exposed birds. These trends supported the anti-detection defense mechanism of concealment. If a bird was concealed in the leaves of a tree, it had no reason to fly away. However, cryptic coloration often allows animals to be in plain view, but still remain hidden. An example of this is Hoffman’s Woodpecker (Melanerpes hoffmanni). Although this bird was exposed and solitary all three times I played raptor calls to it, it displayed no reaction every time. Although there was not much foliage, it was still difficult to spot because it was camouflaged. Therefore, had the woodpecker flown, it would have become visible. The X² table showed a significant value, and the difference in frequency of flights is most apparent in the Exposed X Solitary grid, with 16 total flights (Table 1). This combination supports both of my hypotheses that solitary birds will fly more often than groups and exposed birds will fly more often than hidden birds. If a bird is not using either anti-detection or anti-capture strategies, it makes sense that it will fly if it hears a potential threat. The One-way ANOVA tests showed no relation between bird size and reaction when the entire data set was considered, but showed a trend for flight reactions to be exhibited by larger birds on average when only solitary and exposed birds were considered. The size of the bird was not a reliable predictor of its reaction to the raptor call; the smallest passerines often times only looked around, such as Yellow-Throated Euphonias, Tropical Pewees, and Yellow-Faced Grassquits (Tiaris olivacea) that never flew, while medium-sized birds such as corvids (jays) flew nearly every time. The biggest bird I observed was the Cattle Egret (Casmerodius albus), and it showed no reaction. Raptors may have a tougher time finding the smallest birds, or it may be possible that the raptors do not bother with these smallest birds and instead look for medium sized birds, to be energy efficient. The largest birds may not even be considered prey because of their size. The data show that birds recognize and perceive a threat from raptor calls. The results suggest that birds use the anti-detection strategy of concealment and camouflage, and the anti-capture strategy of flocking. Further research should be done on raptor call recognition on a single species or guild of birds to determine if raptor call recognition is specific on a lower taxonomic level or specific grouping, as previous studies have suggested.

ACKNOWLEDGEMENTS

I would like to thank Alan Masters for his encouragement and guidance on this project, Andrew Rodstrom and Tim Kuhman for their patience and answers to my endless supply of questions, and my parents for supporting me in my desire to explore the tropical forests of Costa Rica.

LITERATURE CITED

Alcock J. 1984. Behavior: An evolutionary approach. Sinauer Associates, Inc., Sunderland, Massachusetts, USA. Buchanan J.B., C.T. Schick, L.A. brennan, and S.G. Herman. 1988. Merlin predation on wintering dunlins: hunting success and dunlin escape tactics. The Wilson Bulletin 100: 108-118. Caldwell G.S. 1986. Predation as a selective force on foraging herons: effects of plumage color anf flocking. The Auk 103: 494-505. Fogden, M. 1993. An Annotated Checklist of the Birds of Monteverde and Penas Blancas. Litografia e Imprenta LIL, San Jose, Costa Rica. Goldman P. 1980. Flocking as a posible predator defense in dark-eyed juncos. The Wilson Bulletin 92: 88-95. Haber W.A., W. Zuchowski and E. Bello. 2000. An Introduction to Cloud Forest Trees: Monteverde, Costa Rica. Mountain Gem Publications, Monteverde, Costa Rica. Hauser M.D. and C. Caffrey. 1994. Anti-predator response to raptor calls in wild crows, Corvus brachyrhynchos hesperis. Animal Behaviour 48: 1469-1471. Holdridge, L.R. 1967. Life Zone Ecology, Revised Edition. Tropical Science Center, San Jose, Costa Rica. Janzen, D.H. 1983. Costa Rican Natural History. The University of Chicago Press, Chicago, Illinois, USA. Lima S.L. and P.A. Bednekoff. Back to the basics of anti-predatory vigilance: can nonvigilant animals detect attack? Animal Behaviour 58: 537-543. Perrins C. 1979. Birds: Their Life, Their Ways, Their World. Reader’s Digest, USA Pettifor R.A. 1990. The effects of avian mobbing on a potential predator, the European kestrel, Falco tinnunculus. Animal Behaviour 39: 821-827. Stiles G.F., and A.F. Skutch. 1989. A Guide to the Birds of Costa Rica. Comstock Publishing Associates, Ithica, N.Y., U.S.A. Terbough, J. 1989. Where Have All the Birds Gone? Princeton University Press, Princeton, N.J., U.S.A. Trail P.W. 1987. Predation and antipredator behavior at Guianan cock-of-the-rock leks. The Auk 104: 496-507. TABLES AND FIGURES

Figure 1. Eighty-seven songbird and 84 raptor calls were played to birds, with the above reactions. Differences were significant for all reactions between songbird and raptor calls.

Figure 2. The Great Blawk-Hawk and Collared Forest-Falcon are residents, while the Semiplumbeous Hawk and Slaty-Backed Forest-Falcon are non-residents; no significant differences. Twenty-one calls were played for each raptor. Figure 3. Average time birds stayed before they flew away (Twenty-three flight reactions total). Raptors 1 & 3 are residents, 2 & 4 are non-residents. Raptor 1 is the Great Blawk-Hawk, 2 is the Semiplumbeous Hawk, 3 is the Collared Forest-Falcon, and 4 is the Slaty-Backed Forest-Falcon.

Figure 4. Fifty-one raptor calls were played for solitary birds, and 33 raptor calls were played for groups, with the indicated reactions (no significant differences between solitary birds and groups). Figure 5. 66 and 18 raptor calls were played for exposed and hidden birds, respectively. No significant differences between exposed and hidden birds for any of the reactions.

Table 1. Conditions under which birds tended to show “fly” reactions in response to raptor calls, X2 = 17.1 Solitary Group Exposed 16 5 Hidden 2 0 APPENDIX

Bird species, size in cm, sequence call # played, # of individuals, exposed (e) or hidden (h), reactions (N – “none”, L – “look”, F – “fly”.

Species Key: A – Grove-Billed Ani, Crotophaga sulciroctris BCM – Blue-Crowned Motmot, Momtus momota BGT – Blue-Grey Tanager, Thraupis episcopus BJ – Brown Jay, Cyanocorax morio BWB – Black-and-White Becard, Pachyramphus albogriseus DCFC – Dusky-Capped Flycatcher, Myiarchus tuberculifer EM – Eastern Meadowlark, Sturnella magna ET – Emerald Toucanet, Aulocorhynchus prasinus GBC – Golden-Browed Chlorophonia, Chlorophonia callophrys GCFC – Gray-Capped Flycatcher, Myiozetetes granadensis GE – Great Egret, Casmerodius albus GK – Great Kiskadee, Pitangus sulphuratus GTG – Great-Tailed Grackle, Quiscalus mexicanus HW – House Wren, Troglodytes aedon HWP – Hoffman’s Woodpecker, Melanerpes hoffmannii ID – Inca Dove, Colombina inca KBT – Keel-Billed Toucan, Ramphastos sulphuratus MIXED – Mixed-species flock MO – Montezuma Oropendola, Psarocolius montezuma MR – Mountain Robin, Turdus plebejus MT – Masked Tityra, Tityra semifasciata NO – Northern Oriole, Icterus g. galbula RCNT – Ruddy-Capped Nightingale-Thrush, Catharus frantzii RCW – Rufous-Capped Warbler, Basileuterus tristriatus SFC – Social Flycatcher, Myiozetetes similus ST – Summer Tanager, Piranga rubra TKB – Tropical Kingbird, Tyrannus melancholicus TP – Tropical Pewee, Contopus cinereus WTMJ – White-Throated Magpie-Jay, Calocitta Formosa WFP – White-Fronted Parrot, Amazona albifrons YFG – Yellow-faced Grazzquit, Tiaris olivacea YTE – Yellow-Throated Euphonia, Euphonia hiruninacea Species Length Seq # # of indivs Exp/Hid Songbird Raptor Time (s) (cm) Rx Rx A 30 3 5 E N N - A 30 1 8 H N L - A 30 1 3 H N N - BCM 39 2 1 E N F 18 BGT 15 1 2 E N L - BGT 15 3 1 E N F 24 BJ 39 2 3 E N F 25 BJ 39 1 3 E N N - BJ 39 2 1 H N F 12 BJ 39 4 1 H N F 12 BWB 14 3 1 E N F 43 DCFC 16.5 1 1 E N F 34 EM 20 4 2 E N F 17 EM 20 3 1 E N F 2 ET 29 3 2 E N L - ET 29 1 1 E N F 18 ET 29 3 1 H L L - ET 29 4 1 H N N - GBC 13 3 9 E N F 36 GCFC 16.5 1 2 E N L - GCFC 16.5 3 2 E N L - GCFC 16.5 3 1 E F - - GCFC 16.5 1 1 E N L - GCFC 16.5 4 1 E N L - GCFC 16.5 4 1 E N L - GE 101 4 1 E N L - GK 23 3 1 E F - - GK 23 2 1 E N F 27 GK 23 1 1 E N L - GK 23 1 1 E N L - GK 23 2 1 E N N - GTG 43 4 1 E N F 1.5 GTG 43 3 1 E N L - GTG 43 2 1 E N N - HW 10 2 1 E N L - HWP 18 1 1 E N N - HWP 18 2 1 E N N - HWP 18 3 1 E N N - ID 20 2 2 E N L - ID 20 4 3 E N N - ID 20 3 2 H N L - ID 20 4 1 E N F 40 ID 20 3 1 H N N - KBT 47 3 1 H N L - MIXED - 4 4 E N N - MO 50 2 2 E N L - MO 50 2 10 E N L - MR 24 1 1 E N L - MR 24 2 1 E N L - MT 21 4 1 E N F 22 NO 18 2 7 E N N - RCNT 16 1 1 H L L - RCW 12.5 1 1 E N F 30 SFC 16 1 3 E N F 26 SFC 16 1 2 E N L - SFC 16 1 2 H N N - SFC 16 3 5 H N N - SFC 16 2 1 E F - - SFC 16 1 1 E N F 32 SFC 16 2 1 E N F 5 SFC 16 4 1 E N F 8 SFC 16 1 1 E N L - SFC 16 1 1 E N L - SFC 16 2 1 E N L - SFC 16 2 1 E L L - SFC 16 3 1 E L L - SHWC 19 4 1 H N N - ST 16.5 2 1 H N L - ST 16.5 2 1 H N L - TKB 21 4 2 E N F 18 TKB 21 2 1 E L F 14 TP 13 3 1 E N L - TP 13 4 1 E N L - TP 13 4 1 E N N - WFMJ 46 3 1 E N F 15 WFP 25 3 2 E N L - WFP 25 4 2 E N N - WFP 25 3 4 H N N - YFG 10 4 3 H N L - YFG 10 2 1 E N L - YFG 10 3 1 E N N - YTE 11 2 4 E N L - YTE 11 4 2 E N L - YTE 11 4 2 E N L - YTE 11 4 3 E N L - YTE 11 3 4 E N N - YTE 11 1 2 H L L -