SPATIAL PATTERNING AND COMPETITION IN AN ASPEN-WHITE PINE SUCCESSIONAL SYSTEM

The spatial patterning of and competition between established large-toothed aspen (Populus grandidentata) and invading white pine (Pinus strobus) were examined in a 65-yr-old forest system at the AuSable Trails Institute of Environmental Studies, Kalkaska County, Michigan. Density, diameter, and distance measures were recorded for tree species on two 40 × 40 m grids containing 64 quadrats each 5 × 5 m. Several indices of dispersion were used to assess pattern and regression analysis of nearest neighbor distance to tree diameter provided an index of competition. The results indicate strong aggregation for both aspen and pine on a large scale, but regularity of pattern for pines in relation to their nearest neighbor. This regularity appears to be due to competition with established aspens and suggests that the spatial patterning of the aspens acts to control the spacing and subsequent development of the invading white pines.     

Social Aggregation in Pea Aphids: Experiment and Random Walk Modeling

From bird flocks to fish schools and ungulate herds to insect swarms, social biological aggregations are found across the natural world. An ongoing challenge in the mathematical modeling of aggregations is to strengthen the connection between models and biological data by quantifying the rules that individuals follow. We model aggregation of the pea aphid, Acyrthosiphon pisum. Specifically, we conduct experiments to track the motion of aphids walking in a featureless circular arena in order to deduce individual-level rules. We observe that each aphid transitions stochastically between a moving and a stationary state. Moving aphids follow a correlated random walk. The probabilities of motion state transitions, as well as the random walk parameters, depend strongly on distance to an aphid''s nearest neighbor. For large nearest neighbor distances, when an aphid is essentially isolated, its motion is ballistic with aphids moving faster, turning less, and being less likely to stop. In contrast, for short nearest neighbor distances, aphids move more slowly, turn more, and are more likely to become stationary; this behavior constitutes an aggregation mechanism. From the experimental data, we estimate the state transition probabilities and correlated random walk parameters as a function of nearest neighbor distance. With the individual-level model established, we assess whether it reproduces the macroscopic patterns of movement at the group level. To do so, we consider three distributions, namely distance to nearest neighbor, angle to nearest neighbor, and percentage of population moving at any given time. For each of these three distributions, we compare our experimental data to the output of numerical simulations of our nearest neighbor model, and of a control model in which aphids do not interact socially. Our stochastic, social nearest neighbor model reproduces salient features of the experimental data that are not captured by the control.     

Influence of environmental factors on the deep and skin temperature in the European bison,Bison bonasus (L.)

• 1.1. In 43 European bison divided into three groups (Group A, 3–8-month-old calves; Group B, 18-month-7-year-old young bison; Group C, 12–24-year-old bison) the rectal, humerus region and abdomen region temperatures were measured.
• 2.2. The experiments were carried out in winter months, from mid-December to mid-March.
• 3.3. The mean rectal temperatures changed from 38.55°C in calves to 38.15°C in the oldest bison.
• 4.4. The mean temperatures of the humerus region changed from 20.69°C in calves to 21.49°C in older bison.
• 5.5. The mean temperatures of the abdomen region changed from 20.79°C in calves to 22.17°C in older bison (Gr. B).
• 6.6. The cluster analysis divided the bison into four groups named hot, warm, cool and cold bison.
• 7.7. Only air temperature measured 2 m above the ground and snow cover influenced the integrated bison temperature. Age, sex and mass as well as some environmental factors had no influence.
• 8.8. Measurements made 1 to nearly 4hr after a bison's death showed a drop in rectal temperature and mostly increases in temperatures of the humerus and abdomen regions.

     

Social Correlates of Variation in Male Calling Behavior in Blanchard's Cricket Frog,Acris crepitans blanchardi

I studied variation in male calling behavior and its social correlates in Blanchard's cricket frog, Acris crepitans blanchardi. Calls were produced in distinct call groups, and they increased in duration and complexity from the beginning to end of a call group. Dominant frequency was the only character of 18 quantified consistently correlated with male snout-vent length. Calls from the beginning of a call group varied independently of calls from the middle and end of a call group, and only calls from the beginning of a call group exhibited significant variation among males, thus relative consistency within males. Other characters varied greatly within individual males. Unlike most other anurans, dominant frequency also exhibited tremendous within-male variation. The relative influence of caller density, local caller density, nearest neighbor distance, and nearest neighbor sound pressure level on variation in male calling behavior was examined. Nearest neighbor distance, mediated through the sound-pressure level of neighbor calls, appeared to have the greatest influence on variation in male calling behavior. The most profound changes in calling behavior occurred during aggressive encounters; males altered their calling behavior in a manner suggesting that they respond to competitors with graded aggressive signals. Furthermore, the structure of the communication system suggests that calls are graded not only in response to the level of social competition, but graded over a call group as well.     

Postrelease Dispersal of Reintroduced Elk (Cervus elaphus) in Ontario,Canada

We studied 2 years of postrelease telemetry data of elk (Cervus elaphus) translocated to their historic range limit in Ontario, Canada and sought to determine if postrelease movements were related to behavior, demography of released animals, or site–specific attributes such as length of holding period. During 1998–2004 we radio‐tracked 341 elk in 10 release groups via ground and aerial telemetry and monitored movement patterns relative to gender, age, and pre‐release holding period (4–112 days). We found that elk that were held for short periods prior to release (4–11 days) moved longer distances than those subject to extended conditioning (17–112 days), suggesting that an extended conditioning period is beneficial from the standpoint of promoting philopatry. When all elk were pooled by sex and age class, male calves remained in closer proximity (8.0 ± 13.2 km) to release sites than adult females (19.1 ± 20.6 km), adult males (19.7 ± 15.1 km), and female calves (14.4 ± 20.4 km). Most calves dispersed in a southeasterly direction whereas adults tended to travel southwest. Our results reveal that elk movement characteristics are influenced by factors such as release protocol and group demographics; these findings provide further insight regarding appropriate release methods for restoring natural populations near their historical range limit.     

Pen size and parity effects on maternal behaviour of Small-Tail Han sheep

The aim of this experiment was to study the effects of pen size and parity on maternal behaviour of twin-bearing Small-Tail Han ewes. A total of 24 ewes were allocated to a 2×2 design (six per pen), with parity (primiparous or multiparous) and pen size (large: 6.0×3.0 m; small: 6.0×1.5 m) as main effects at Linyi University, Shandong Province, China. Behaviour was observed from after parturition until weaning. All ewes were observed for 6 h every 5 days from 0700 to1000 h and from 1400 to 1700 h. Continuous focal animal sampling was used to quantify the duration of maternal behaviours: sucking, grooming and following as well as the frequency of udder accepting, udder refusing and low-pitched bleating. Oestradiol and cortisol concentrations in the faeces (collected in the morning every 5 days) were detected using EIA kits. All lambs were weighed 24 h after parturition and again at weaning at 35 days of age. The small pen size significantly reduced following (P<0.005), grooming (P<0.001) and suckling durations (P<0.05), as well as the frequency of udder refusals (P<0.001). However, there was a significant interaction with ewe parity, with decreased grooming and suckling in the small pen largely seen in the multiparous ewes (P<0.001). Independent of pen size, multiparous ewes accepted more sucking attempts by their lambs (P<0.05) and made more low-pitched bleats than primiparous ewes (P<0.001). Multiparous ewes had higher faecal oestradiol concentrations than primiparous ewes (P<0.001), and ewes in small pens had higher faecal cortisol levels compared with ewes in larger pens (P<0.001). As lambs increased in age, the duration of maternal grooming, following and suckling as well as frequency of udder acceptance and low-pitched bleating all declined, and the frequency of udder refusing increased (P<0.001 for all). Ewe parity, but not pen size, affected lamb weight gain during the period of observation (P<0.001). This is the first study to show that pen size, interacting with parity, can affect the expression of maternal behaviour in sheep during lactation. The study is also the first to report on the maternal behaviour of Chinese native sheep breeds (Small-Tail Han sheep), with implications for the production of sheep in China.     

Factors Affecting Group Spread within Wild Mixed-Species Troops of Saddleback and Mustached Tamarins

We examined group spread and interindividual spacing within wild mixed-species troops of saddleback (Saguinus fuscicollis) and mustached (Saguinus mystax) tamarins. Mustached tamarin groups were spread over larger areas than those of saddleback tamarins. Group size and behavior affected group spread and interspecific proximity: larger groups of both species were dispersed over greater areas, and the larger troop had the lowest degree of interspecific proximity. Behavior also affected group spread and interspecific proximity: when traveling individuals were spread over a larger area, and the distance between heterospecifics was greater than when stationary. We examined spatial proximity using data on the distance from nearest neighbor. Overall, distance to nearest neighbor was not affected by group size. Based on specific behaviors, foraging saddleback tamarins were significantly nearer conspecifics than to mustached tamarins. Tamarins have polymorphic color vision, and trichromats—having 3 types of visual pigment—versus dichromatic individuals—with two types of pigment—may be better at perceiving yellow or russet colored predators and conspecifics than their dichromatic counterparts are. Color vision status affected spatial positioning, with vigilant trichromats being further from their neighbors than their dichromatic conspecifics were. We discuss the findings with respect to the ecology of the species. Specifically, interspecific differences in group spread and spatial proximity are related to differences in the supports used, and the effect of troop size on interspecific proximity is related to increased resource competition. The finding that trichromats are further from their neighbors represents the first example of a behavioral correlate of color vision ability in a wild species with polymorphic color vision, and is explained through the perception of predation risk.     

种群分布格局的多尺度分析

种群分布格局的分析对于了解种群空间分布规律以及种内与种间关系具有重要的意义。最近邻体分析方法 (Nearestneighboranalysis, NNA) 作为种群空间分布格局的重要分析方法, 仅局限于种群格局的单尺度分析。改进NNA方法以应用于种群格局的多尺度分析, 将有助于解决种群格局的尺度依赖性。该文在前人研究的基础上提出扩展最近邻体分析方法 (Extendednearestneighboranalysis, ENNA), 也即在传统Clark_Evans指数公式的基础上增加一个距离尺度参数d (m), 并定义其所对应的Clark_Evans 指数CE (d) 的计算公式及其相应的显著性检验计算公式 (u (d) ) 分别为 :CE (d) =rdA/rdE= (1Nd∑Ndi=1 rdi) / (0.5Ad/Nd+0.0 5 14Pd/Nd+0.0 4 1Pd/Nd3 /2 ) 和u (d) = (rdA-rdE) /σd, 在距离尺度d (m) 范围内, 参数rdA指样地内各个体与其最近邻体间距离的平均值 (m) 、rdE指相同环境中个体呈随机状态时最近邻体距离的平均值 (m) 、Nd 为样地内个体总数、rdi为第i个个体与其最近邻体间的距离 (m) 、Ad 为样地面积 (m2 ) 、Pd 为样地周长 (m) 和σd 代表标准差。ENNA尺度变换采用与分形理论中计算沙盒维数相类似的过程, 而格局类型判断的标准与传统最近邻体分析方法相同。传统最近邻体分析结果是EN NA中距离尺度d取最大值dmax时的一个特例。以广东省黑石顶自然保护区针阔叶混交林中的马尾松 (Pinusmas soniana) 、黄牛奶树 (Symplocoslaurina) 、水栗 (Castanopsisnigrescens) 、鼠刺 (Iteachinensis) 和桃金娘 (Rhodomyrtustomentosa) 等 5个代表性种群为例, 在地理信息系统软件ArcViewGIS技术平台上进行的实例研究显示, 5个种群均表现出不同程度的尺度相关性。由此表明, 该文提出的新方法ENNA能够检测出种群空间分布格局的尺度依赖性, 获得关于种群空间分布格局的多尺度信息, 是进行种群空间格局多尺度分析的有效方法。     

湖北九宫山成熟常绿落叶阔叶混交林3种优势种的邻域效应研究

以最近邻株距离统计研究了湖北九宫山成熟常绿落叶阔叶混交林3种优势种青冈栎(Cyclobalanopsis glauca)、甜槠(Castanopsis eyrei)和短柄枹栎(Quercus serrata var.brevipetiolata)的邻域效应,进而探讨了局域尺度上的竞争/帮促关系与物种共存格局.结果显示,青冈栎、甜槠和短柄枹栎的最近邻株距离不存在下限;青冈栎的胸径大小与其种内最近邻株距离存在正相关(p=0.029),甜槠的胸径大小与种内、种间最近邻株距离均无相关性(p≥0.360),而短柄枹栎的胸径大小与种间最近邻株距离为负相关(p=0.040);3树种中任意一个的种内与种间最近邻株距离都没有显著差异(p≥0.122).这些结果表明,九宫山成熟常绿落叶阔叶混交林内邻域尺度上的竞争排斥并未充分激化,但是青冈栎的种内竞争仍然在起作用,而甜槠的竞争效应不明显,短柄枹栎则依赖种间帮促甚于相互竞争.同种和异种之间的排斥效果没有差异表明局域尺度的物种共存格局可能出于随机过程而非负密度制约过程.     

湖北九宫山成熟常绿落叶阔叶混交林 3种优势种的邻域效应研究

以最近邻株距离统计研究了湖北九宫山成熟常绿落叶阔叶混交林3种优势种青冈栎(Cyclobalanopsis glauca)、甜槠(Castanopsis eyrei)和短柄枹栎(Quercus serrata var.brevipetiolata)的邻域效应,进而探讨了局域尺度上的竞争/帮促关系与物种共存格局。结果显示,青冈栎、甜槠和短柄枹栎的最近邻株距离不存在下限;青冈栎的胸径大小与其种内最近邻株距离存在正相关(p= 0.029),甜槠的胸径大小与种内、种间最近邻株距离均无相关性(p≥ 0.360),而短柄枹栎的胸径大小与种间最近邻株距离为负相关(p= 0.040);3树种中任意一个的种内与种间最近邻株距离都没有显著差异(p≥ 0.122)。这些结果表明,九宫山成熟常绿落叶阔叶混交林内邻域尺度上的竞争排斥并未充分激化,但是青冈栎的种内竞争仍然在起作用,而甜槠的竞争效应不明显,短柄枹栎则依赖种间帮促甚于相互竞争。同种和异种之间的排斥效果没有差异表明局域尺度的物种共存格局可能出于随机过程而非负密度制约过程。     

Arena behaviour of ewes selected for superior mothering ability differs from that of unselected ewes

As part of a study searching for criteria for the genetic improvement of maternal ability, behavioural parameters were measured in ewes from two flocks, one selected for maternal ability (Fertility Flock), the other unselected (Random Flock). Behaviour was measured in an open-field test prior to mating in 1990 and in an arena test prior to mating in 1990 and 1991, and prior to lambing in 1991.The open-field test consisted of placing the ewes in an enclosed field measuring 16 m × 16 m for a period of 5 min. The field was divided into 16 squares each measuring 4 m × 4 m. The following measures were made: number of squares entered, whether or not the ewe bleated, urinated or defaecated and the number of bleats.The arena test consisted of placing the ewes in an enclosed arena measuring 3 m × 13 m; at one end of the arena there was a pen of sheep, in front of which sat a person. The floor of the arena was marked into 39 squares each measuring 1 m × 1 m. The test lasted 10 min and the following parameters were derived by recording the square in which the animal was located every 15 s; minimum and maximum distance from the person, mean distance from the person and distance travelled. The number of bleats was also recorded for each ewe, as well as whether or not she bleated, urinated or defaecated.There were no differences between the two flocks in any of the parameters measured in the open-field test. However, on all three occasions, there were differences between the two flocks in the total distance travelled and the number of bleats. The coefficients of repeatability of these two parameters, as measured by the intra-class coefficient of correlation, were 0.48 and 0.57 respectively.The results indicate that the arena test measures differences in the behaviour of ewes that differ in maternal ability and may therefore be of use in identifying ewes of superior mothering ability.     

个体大小对草鱼耐高温能力及升温过程中群体行为的影响

为考察个体大小对升温过程中草鱼的耐高温能力和群体行为的影响, 研究以草鱼(Ctenopharyngodon idellus)幼鱼为实验对象, 结果显示: 大个体群体的个体游泳速度明显大于小个体群体(P=0.039), 个体间距离和最近邻距离显著高于小个体群体(P<0.001); 大个体鱼群的极性与小个体群体并无差异。虽然大个体群体的致死高温与小个体群体无差异, 但前者的临界高温显著大于小个体群体(P<0.001)。升温过程会使鱼群个体游泳速度变快, 个体间距离和最近邻距离也相应增加, 但游泳速度同步性和群体极性维持不变。研究表明: 个体大小的增加可增强草鱼幼鱼对环境水温的耐高温能力; 升温不会影响不同个体大小草鱼幼鱼群体的协调性, 但会降低草鱼幼鱼的群体凝聚力。     

The formation of preferential relationships at early age in cattle

Calves can develop long-lasting social relationships with peers. We examined the strength of the relationships between calves according to the time they had been together. Twenty-four female dairy calves were assigned to six groups of four animals (Type-1 partners) at 0.5 month of age. At 3.5 months of age, they were mixed with other calves (Type-2 partners) to form groups of 14. Type-3 partners were calves added to the experimental groups after 5.25 months. The calves stayed together until 1.5 years of age. Social preferences between the three partner types were examined in a Y-maze, and the position and activity of animals in the barn and pasture were followed in three periods. Behavioural synchrony, distance between animals, proximity and nearest neighbour were analysed. The calves more frequently butted Type-3 than Type-1 partners in the Y-maze (P < 0.05). They spent more time in proximity to Type-1 partners, and these were more often the nearest neighbours than other partners (P < 0.001). Synchrony and distance between animals were greater at pasture than in the barn (P < 0.01). Calves seem to form preferential relationships before 3.5 months of age. Keeping cattle together from an early age seems beneficial for them.     

Social dominance relationships and spacing behavior of swine

Dominance ranks were determined within each of eight pens of domestic swine, Susscrofa. Data on spacing and orientation of subjects were obtained from 16 mm time-lapse films and 35 mm overhead photographs with the aid of a digitizer assisted FORTRAN program, DISANGL 2. Dominance orders among subjects were relatively linear and stable. No pattern in pen location preferences (fixed space) was found but there were relationships between dominance rank and portable spacing patterns of the subjects. Distance to a nearest neighbor was the most important variable affecting the orientation of the swine, and the most dominant animal had a lesser tendency to face away from nearest neighbors at any particular distance and a greater tendency to space further from nearest neighbors than was the case for subordinates. The results of this study support the notion that social dominance provides a means by which a possible limited resource, such as personal space, may be differentially allocated to group members.     

Dosage response evaluation of luprostiol administered to pregnant sows

Two trials were completed to investigate the effects of luprostiol in swine. The first trial was to evaluate parturition induced by various dosages of luprostiol compared with those of lutalyse or vehicle. Sows were assigned by random allotment to one of the following treatments on Day 112 of gestation: Group A, control (0 mg luprostiol); Group B (1.88 mg luprostiol); Group C (3.75 mg luprostiol); Group D (7.5 mg luprostiol); Group E (15 mg luprostiol); Group F (10 mg lutalyse). All prostaglandin-treated groups farrowed earlier than the controls (P<0.05), with Groups D (26.3 h), E (31.0 h) and F (25.8 h) having the shortest treatment-to-first-pig intervals, and Groups A (76.0 h), B (54.4 h) and C (40.0 h) having the longest intervals. Luprostiol-treated sows had the shortest farrowing time (P<0.05; range = 3.2 to 3.9 h). Significant differences were found for the time (min) between births: Group A (32.1), Group B (28.4), Group F (35.5) took longer than Group C (20.2), Group D (21.0) and Group E (21.6). In a second trial, 20 crossbred pregnant sows received either vehicle or luprostiol (7.5 mg) on Day 112 of gestation. Progesterone concentrations declined rapidly (P<0.05) in luprotiol treated females but were unchanged in control females during the 24-h collection period. The results of these trials show 7.5 mg luprostiol to be the most effective dose for inducing farrowing.     

Sucking behaviour of dairy calves fed milk ad libitum by bucket or teat

The milk intake and sucking behaviour of Dutch and Holstein-Friesian crossbred calves fed milk replacer ad libitum by either the bucket method (n=8) or an artificial teat (n=6) were compared. The animals were observed for 3 weeks from the age of 2 weeks, penned individually and all provided with a “dummy” (artificial) teat near the milk source within the pen.

The teat-fed calves ingested significantly more milk than the bucket-fed calves (11.9 vs. 8.0 kg/day; P<0.05), and this intake took much longer (44.2 vs. 17.7 min/day; P<0.05). In both treatment groups, milk intake was organized in “meals”. The meal criterion, separating the withinmeal and between-meal non-feeding intervals, was set at 5 min. Frequency of meals and daily total meal duration did not differ significantly. Meals occurred rather randomly throughout a 24-h period.

On average, the dummy teat was used for 13 min per day by bucket-fed calves, but for only 1 min by teat-fed calves (P<0.05). Sucking of the dummy teat was largely clustered within the meal periods.

It is concluded that in the young calf a need for sucking exists independently of milk satiation. However, the level of satiation depends on whether the calf drinks or sucks the milk. Nutritive sucking is clearly more reinforcing than non-nutritive sucking.     

Estrous behavior following prostaglandin F2α injection in Zebu cattle under continuous observation

A study was conducted to improve understanding of the behavioral aspects of Zebu cattle in estrus in the tropics. Sixteen of 26 cows in group A and 21 of 34 in group B were found to have a corpus luteum and were injected with 30 mg of prostaglandin F2α. In group B, a bull of proven libido was placed with the cows. Animals not injected were considered to be in proestrus and therefore left in the groups. Recorded sexual activity included licking and smelling the genital area, butting, following, and mounting or attemping to mount, both actively and passively. Animals were observed continuously for 100 hs. following injection. Only 56% of the injected animals in Group A and 62% in group B were found in estrus. The average number of mounts observed was 15.9 per cow during the estrous period which lasted 14.4 h. ± 5.1 (mean ± SD) for 12 cows in group A. In group B, cows mounted 13.9 times each in a period of 12.6 h. ± 5.5 (mean ± SD) for 17 animals. In both groups, the 3-hour period during which most mounting activity was recorded was from 0600 to 0900 (25% and 33% for groups A and B, respectively). However, 63% of all mounts were observed at night (1800 to 0600 hrs) in group A as were 53% in group B. No sexual activity was observed for any cow during the first 40 h. of observation. Activity started with constant butting that preceded mounting behavior. This pattern was better defined in group A than in group B where the bull was present. Although all activities were observed at some stage during the observation period, no clear relationship between mounting and other activities could be established.     

PLOIDAL CHANGES IN CLONAL CULTURES OF SPIROGYRA COMMUNIS AND IMPLICATIONS FOR SPECIES DEFINITION

A clonal culture of Spirogyra filaments of initially uniform width produced filaments of three additional significantly different widths. Group I filaments of the original clone were 30.9 ± 0.7 μm wide (mean ± SD, N = 50). Group I filaments produced Group II filaments (22.0 ± 1.1 μm) through vegetative growth and sexual reproduction. Zygospores from homothallic Group I filaments produced germlings representative of Groups I and II; zygospores from homothallic Group II filaments produced germlings representative of Group II only. Germlings of Groups III (27.7 ± 1.0 μm) and IV (44.9 ± 0.8 μm) were produced in the cross of I × II. Viable zygospores from homothallic Group III filaments were obtained. Cells of Group IV filaments were initially binucleate and did not conjugate. Of the six intergroup crosses possible, four resulted in conjugation-tube formation only; two crosses yielded zygospores (I × II and III × IV). Germlings from the successful cross of Groups III and IV produced filaments of all four groups. Chromosome counts were: Group I (24), Group II (12), Group III (18), and Group IV (24, one nucleus). Relative nuclear fluorescence values of mithramycin-stained DNA were (mean ± SD, N ≥ 30): Group I (11.1 ± 1.4), Group II (5.7 ± 0.7), Group III (8.8 ± 1.3), and Group IV (10.0 ± 0.9, one nucleus). Cytologically, Group II appears to be a diploid (2x), Group I a tetraploid (4x), and Group III a triploid (3x). Systematically, Groups I, II, and III key out to Spirogyra singularis, S. communis, and S. fragilis, respectively, using Transeau's mongraph of the family Zygnemataceae. These species are interpreted to represent a species complex of S. communis (whose name has priority) with the ancestral haploid (x = 6) missing.     

Detecting spatial aggregation from distance sampling: a probability distribution model of nearest neighbor distance

Spatial point pattern is an important tool for describing the spatial distribution of species in ecology. Negative binomial distribution (NBD) is widely used to model spatial aggregation. In this paper, we derive the probability distribution model of event-to-event nearest neighbor distance (distance from a focal individual to its n-th nearest individual). Compared with the probability distribution model of point-to-event nearest neighbor distance (distance from a randomly distributed sampling point to the n-th nearest individual), the new probability distribution model is more flexible. We propose that spatial aggregation can be detected by fitting this probability distribution model to event-to-event nearest neighbor distances. The performance is evaluated using both simulated and empirical spatial point patterns.     

Aggression and nest spacing in single and mixed species groups of seabirds

When heterospecific seabirds are part of a nesting colony, there may be less opportunity for conspecifics to come in direct contact with each other, resulting in lower intraspecific aggressiveness. To determine if individuals spend less time in aggressive behavior when nesting in conspecific rather than heterospecific groups, we compared the behavior of black skimmers (Rhynchops niger) nesting with gull-billed terns (Sterna nilotica) in three mixed species subcolonies to those of black skimmers in three single species subcolonies. In contrast to our predictions, black skimmers spent significantly less time in aggressive behaviors when nesting in single species subcolonies than when nesting with heterospecifics. Although skimmers in mixed species subcolonies tended to have more aggressive interactions with skimmers than terns, this may be a function of subcolony composition; the proportions of aggressive interactions with conspecifics were similar to the proportions of conspecifics in each subcolony. However, within the mixed species subcolonies, skimmers that nested nearer to terns were involved in aggressive interactions significantly less than skimmers that nested closer to conspecifics. Also, skimmers nested closer to their nearest neighbor when it was a gull-billed tern than when it was another skimmer. Regardless of which species they nested closest to, skimmers were more aggressive towards other skimmers than to terns within the mixed species subcolonies. Distance to nearest neighbor's nest did not differ significantly between the colony types, and did not seem to influence the duration of aggressive activity in the single species subcolonies. In the mixed species subcolonies, however, the time spent in aggressive behavior increased as the distance to nearest neighbor increased. It appears that of the several benefits that have been proposed of mixed species colonies, reduced time spent in conspecific aggression is not among them. However, within a mixed species colony, an individual can reduce time spent in aggressive interactions by nesting near heterospecifics. Received: 16 September 1996 / Accepted: 24 February 1997