Modeling the flocking propensity of passerine birds in two Neotropical habitats

We examined the importance of mixed-species flock abundance, individual bird home range size, foraging height, and foraging patch characteristics in predicting the propensity for five Neotropical passerine bird species (Slaty Antwren, Myrmotherula schisticolor; Golden-crowned Warbler, Basileuterus culicivorus; Slate-throated Redstart, Myioborus miniatus; Wilson’s Warbler, Wilsonia pusilla; and Black-and-white Warbler, Mniotilta varia) to forage within flocks, rather than solitarily. We used study plots in primary mid-elevation forest and in shade coffee fields in western Panama. We expected that all species would spend as much time as possible flocking, but that the social and environmental factors listed above would limit compatibility between flock movements and individual bird movements, explaining variability in flocking propensity both within and among species. Flocking propensity was well predicted by home range size and flock abundance together, for four of the five species. While flock abundance was uniform across plots, home range sizes varied among species and plots, so that home range size appeared to be the principle factor limiting flocking propensity. Estimates of flock abundance were still required, however, for calculating flocking propensity values. Foraging height and patch characteristics slightly improved predictive ability for the remaining species, M. miniatus. In general, individual birds tended to join flocks whenever one was available inside their home range, regardless of a flock’s specific location within the home range. Flocking propensities of individual species were lower in shade coffee fields than in forests, and probably vary across landscapes with variations in habitat. This variability affects the stability and species composition of flocks, and may affect survival rates of individual species. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Flamingo breeding: The role of group displays

Group displays in flamingos have been presumed to play a role in stimulating synchronous nesting and in facilitating pair formation. This study compares the group displays and breeding success of a captive flock of Caribbean flamingos (Phoenicopterus ruber ruber) at the National Zoological Park between 2 years: the frequency and synchrony of group displays were measured for a flock of 17 in 1988 and then again in 1989 after flock size was increased to 21. In 1989 the rate of occurrence of display activity increased 48%, the synchrony of group displays increased 100%, the frequency of mounts and copulations almost doubled, and for the first time in the flock's history two fertile eggs were produced. The use of sprinklers to simulate rain had no effect on the group displays. The amount of naturally occurring rainfall in 1989 was almost twice that in 1988. The increased frequency and synchrony of group displays could be attributed to increased flock size, change in sex ratio, addition of strange individuals, or increased rainfall. This study, however, provides evidence for a relationship between behavioral stimulation from group displays and components of breeding success in flamingos.     

Effects of snow cover on the social and foraging behavior of the great tit Parus major

In the Joetsu region of central Honshu, Japan, snow lies on the ground 2–3 m deep from January to February every year. To test the effects of snow cover on the social and foraging behaviors of the great tit (Parus major Linnaeus) in the region, the following parameters were compared between the pre-snowy period (November and December) and the snowy period (January and February) at the individual level: population size, monospecific flock size and the stability of its membership, home range size, attendance rate with mixed-species flocks, and foraging height. Great tits lived alone or in pairs, but often joined mixed-species flocks. Throughout the study period, individuals exhibited strong site fidelity, which resulted in a stable population size. Neither the size of a monospecific flock nor its membership was affected by snow cover. The home range expanded when birds joined mixed-species flocks during both periods. Birds more frequently joined mixed-species flocks during the snowy period, and the size of mixed-species flocks was significantly larger than during the pre-snowy period. During the snowy period, birds shifted their foraging position from the ground to the upper parts of trees. They also used the upper parts of trees when they joined mixed-species flocks. These results suggest that the intraspecific sociality of great tits is relatively insensitive to snow cover, but that the home range size and foraging positions are affected by joining mixed-species flocks, rather than by snow cover.     

滨海人工湿地越冬鸟类混合群结构及角色

2013年12月至2014年2月冬季,采用样点法对广西防城港市防城区江山乡新基村虾塘的越冬鸟类混合群行为进行观察研究。鸟类混合群在深水虾塘与浅水虾塘生境持续的平均时间分别为(51.6±33.6)min/群与(31.9±13.3)min/群。平均水深15 cm的深水虾塘与平均水深5 cm的浅水虾塘中均分布有11种鸟类,深水虾塘的主要水鸟为体型中等的鹤鹬(Tringa erythropus)、青脚鹬(T.nebularia)、泽鹬(T.stagnatilis),浅水虾塘的主要水鸟是小型个体的金眶鸻(Charadrius dubius)、青脚滨鹬(Calidris temminckii)。深水虾塘鸟类混合群的平均物种数及个体数分别为(5.48±1.60)种/群和(18.75±11.67)只/群,浅水虾塘鸟类混合群则分别为(3.93±1.14)种/群和(11.65±5.12)只/群。深水虾塘的核心种鹤鹬以及跟随种青脚鹬、泽鹬在浅水虾塘属于加入种;而浅水虾塘的核心种金眶鸻以及跟随种青脚滨鹬在深水虾塘却属于加入种。研究结果表明,不同类型生境的鸟类混合群结构及其核心种和跟随种均存在明显差异。     

Description of a new species from Laguna Chichancanab,Yucatan, Mexico: <Emphasis Type="Italic">Cyprinodon suavium</Emphasis> (Pisces: Cyprinodontidae)

Cyprinodon suavium is a new species that belongs to the endemic species flock from Laguna Chichancanab, Yucatan, Mexico, which is proposed to have evolved by sympatric speciation in the lake during the last 8000years. C. suaviumis distinguished from all other known Cyprinodon species by a flattened and concave inter- and postorbital skull roof and a terminal mouth with distinctive thickened lips. The short gut length and dietary items found in the gut indicate that C. suavium is one of the carnivorous members of the flock.     

Variation in social rank acquisition influences lifetime reproductive success in black-capped chickadees

Dominance relationships structure many animal societies, yet the process of rank attainment is poorly understood. We investigated acquisition of social dominance in winter flocks and its fitness consequences in male black-capped chickadees (Poecile atricapillus) over a 10-year period. Age was the best predictor of rank, and paired comparisons showed high-ranked males to be older than their low-ranked flock-mates. When controlling for age, morphological variables did not predict male social rank, but high-ranked males were heavier, had lower fat scores and were in leaner condition than low-ranked males. Males that survived between years tended to increase in rank over time; however, the rate of rank advancement varied individually. Rank reversals between familiar contestants were rare, and changes in male social rank were associated with changes in flock membership. Average lifetime reproductive success (LRS) of males and females was variable and best predicted by lifespan. Male rank history also influenced realized reproductive success. Birds with higher average rank over their lifespan were more likely to reproduce successfully. However, among successful birds, average rank did not significantly predict LRS. Thus, birds that lived longer and attained high social rank earlier had higher fitness, but this effect was not manifested as fine-scale differences among successful individuals. Taken together, these findings demonstrate the importance of social factors influencing individual fitness.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 85–95.     

Managing the social environments of flamingos for reproductive success

Recent research has shown that large flamingo flocks are more likely to experience reproductive success than small flocks and that there is a positive relationship between behavioral stimulation from group displays and reproductive success. This study compares the group displays and reproductive success in a captive flock of Caribbean flamingos over 6 years at the National Zoological Park. Birds were added to the flock in two different years. In each case, the flock produced fertile eggs during the breeding season following the addition of new birds. Furthermore, the addition of new birds in 1988, and then again in 1991, significantly increased group display activity over the previous year when the flock size was lower. That the addition of birds stimulated group display activity suggests an alternative management strategy for promoting reproduction in captivity: separate birds in a flock and then reunite them prior to the breeding season. © 1994 Wiley-Liss, Inc.     

Reproductive segregation among the Barbus intermedius complex of Lake Tana,Ethiopia. An example of intralacustrine speciation?

Reproductive segregation among the large barbs Barbus intermedius complex, Cyprinidae , of Lake Tana, Ethiopia, was investigated in the mouths and upstream reaches of rivers tributary to the lake, during the spawning seasons of 1993 and 1994. The percentage of running fish of four Barbus morphotypes [Acute (Ac), Bigmouth big-eye (Be), Intermedius (In), and Shorthead (Sh)] peaks with the highest water level of the lake. Large differences are apparent in the minimum fork length of running fish among morphotypes [c. 12 cm for male Sh and 28 cm for male Bigmouth small-eye (Se)]. Significant differences occur in temporal, but especially in spatial patterns of the relative abundance of running morphotypes, when the lake is compared with the four major river mouths and their upstream reaches. These different relative abundances are clearest among Ac, Be, In, Sh and Troutlike (Tr). Also, the absence from the rivers of some morphotypes that are usually abundant in the lake itself [Barbel (Ba), Dark (Da), Lip (Li), and Zurki (Zu)], and the abundant presence in the rivers of Tr, rare in the lake, strengthen the hypothesis that spatial and/or temporal spawning segregation is present among the morphotypes. There are indications that Bighead (Bh) and Zu spawn in the lake itself, while other morphotypes were only found running in or near rivers. Combined with the morphological, ecological, and genetic differences, these data suggest the existence of several Barbus species, forming a unique species flock, which probably evolved within the lake. The presence of several, instead of one Barbus species complicates management of sustainable fisheries and protection of the lake's biodiversity. Present results show the importance of the rivers for Barbus reproduction and stress the urgent need for careful management of fishing activities in spawning times and areas.     

Phylogenetic relationships of species of Crenicichla (Teleostei: Cichlidae) from southern South America based on the mitochondrial cytochrome b gene

Phylogenetic analysis using Bayesian inference, likelihood and parsimony methods was conducted on 60 complete mitochondrial cytochrome b sequences from 21 species of Crenicichla, including all species known from Uruguay (Crenicichla celidochilus, Crenicichla lepidota Crenicichla minuano, Crenicichla missioneira, Crenicichla punctata, Crenicichla scottii, Crenicichla vittata), Crenicichla compressiceps, Crenicichla empheres, Crenicichla geayi, Crenicichla iguassuensis, Crenicichla macrophthalma, Crenicichla menezesi, Crenicichla notophthalmus, Crenicichla regani, Crenicichla cf. regani, Crenicichla semifasciata, Crenicichla sveni, Crenicichla tendybaguassu, two unidentified species, and also two species of Teleocichla. Bayesian analysis resulted in a trichotomy with three major groups: (1) The C. missioneira species group (C. celidochilus, C. empheres, C. minuano, C. missioneira, C. tendybaguassu, and an undescribed species analyzed); (2) a group of southern species (C. iguassuensis, C. punctata, C. scottii, C. vittata); and (3) a rather heterogeneous group comprising the type species C. macrophthalma, members of the Crenicichla reticulata species group (C. geayi, C. semifasciata), members of the Crenicichla wallacii species group (C. compressiceps, C. notophthalmus, C. regani, C. cf. regani), members of the Crenicichla saxatilis species group (C. lepidota, C. menezesi, C. sveni, C. sp.), and two species of Teleocichla. Parsimony jackknifing resulted in a quadritomy with: (1) C. macrophthalma, (2) Teleocichla, (3) the saxatilis + wallacii group species, and (4) the rest, which include C. geayi and C. semifasciata as sister group to a dichotomy with the C. missioneira group and the remaining southern species. The sequence variation within the C. missioneira group is remarkably minor despite considerable morphological differences, supporting the conclusion that it forms an endemic species flock in the Uruguay River basin. Previously proposed species groups within the speciose genus Crenicichla (more than 90 species known) are partly corroborated. However, C. celidochilus was not previously associated with the C. missioneira species group, and C. vittata has not previously been associated with C. scottii, C. iguassuensis, or C. punctata. Crenicichla lepidota, C. sveni, C. menezesi and C. sp. represent the C. saxatilis group. Species of small size, representing the C. wallacii species group and Teleocichla are characterized by very long branches, and the position of Teleocichla differed considerably between the Bayesian and parsimony trees. This finding does not invalidate Teleocichla but rather suggests that the several monophyletic major clades within Crenicichla may need nominal recognition. A putative hybrid specimen with a morphology combining components from C. vittata and C. scottii, but with a cytochrome b sequence from C. scottii was found in a sample from the Rio Quaraí/Cuareim. Another putative hybrid specimen with a unique morphology but a cytochrome b sequence agreeing with C. scottii was found in a sample from Maldonado, but no other Crenicichla species than C. scottii is known from that locality.     

Lake Tana large barbs: phenetics, growth and diversification

Among the Lake Tana large barbs of about 10 cm SL only representatives of the' acute'morphotype can be distinguished, in the size range 10–20 cm SL' bigmouth big eye' can be identified also. As for the rest, very few individuals can be confidently affiliated with a particular morphotype most of them looking like' intermedius'. Even within the range of 20–30 cm SL some individuals are still difficult to identify. Principal component analysis of cranial characters revealed discrete groups of morphotypes. Differences in both external and cranial characters of the morphotypes result from divergence which is most pronounced when fish are 4–5 years old and 20–25 cm SL. This divergence cannot be related exclusively with differences in the growth rates of individuals representing different morphotypes. Differences in food composition between the morphotypes probably increase in parallel with their morphological divergence. Differences between the morphotypes in the lateral line (11) and the gill rakers (Sb) counts were revealed using ANOVA. Comparison of the Lake Tana Barbus complex of forms with that previously known from Lake Lanao (Philippines) suggests that in both lakes the different forms arose sympatrically but that sympatric speciation in Lake Lanao has advanced further than in Lake Tana.     

Genetic divergence among morphotypes of Lake Tana (Ethiopia) barbs

The genus Barbus is a paraphyletic assemblage that includes three different ploidy levels: diploid, tetraploid and hexaploid. 'B.' intermedins , which inhabits Lake Tana (Ethiopia), is a hexaploid taxon that forms a 'species flock' consisting of at least 14 morphotypes. Thirty-one presumptive allozymic loci were analysed in a large sample of 'B.' intermedius. No diagnostic loci were observed between the morphotypes, but substantial differences in allele frequencies were found. One morphotype, Acute, differs significantly from the others. The presence of this genetic differentiation within the lake strongly suggests that the morphotypes are diverging lineages. Because of some parallel characteristics with cichlid species flocks, we put forward the hypothesis that the morphotypes of barbs in Lake Tana represent a species flock that originated via sympatric speciation with more rapid morphological than molecular differentiation. If the differentiation and the reproductive isolation are real, it is probable that the morphotypes already are or will soon reach the status of species.     

黑嘴端凤头燕鸥北方群体初析

黑嘴端凤头燕鸥(Thalasseus bernsteini),今又称中华凤头燕鸥,因其群体数量殊微自鸟种受胁状态定量评估以来一直被视为极危物种。尽管近些年来只观测报道了该种在台湾海峡和大舟山群岛的两个群体,作者2010年撰文指出应视其北方群体依然存在。2016年8月8日,韩国政府发布公报称,在韩国西南部的近海无人岛屿上观察记录到有黑嘴端凤头燕鸥繁殖,同年自8月初至9月上旬,在中国江苏如东和山东胶州湾相继发现记录到黑嘴端凤头燕鸥成鸟引领当年幼鸟并成鸟结小群停歇的多起实例。据此,对黑嘴端凤头燕鸥北方群体的存在应再无争议且对该种繁殖分布态势之拼图板的认知亦初显完整。但诘问依然是:就黑嘴端凤头燕鸥群体总量而言,被人类所知晓的,或者换言之,处在人类监控或掌控之下的,其比例究竟如何?     

Nuclear gene variation and molecular dating of the cichlid species flock of Lake Malawi

The cichlid fishes of Lake Malawi are famously diverse. However, evolutionary studies have been difficult because of their recent and uncertain phylogenetic history. Portions of 12 nuclear loci were sequenced in nine rock-dwelling species (mbuna) and three representatives of pelagic nonmbuna species. In contrast to the pattern of variation at mitochondrial genes, which do provide phylogenetic resolution at the level of mbuna versus nonmbuna, and among some genera, the nuclear loci were virtually devoid of phylogenetic signal. Only a small minority of variable positions were phylogenetically informative, and no phylogenetic branches are supported by more than one site. From the nuclear gene perspective the Malawian radiation appears to be a star phylogeny, as if the founding of the lake was accompanied by a partial bottleneck. The pattern is different from that found in Lake Victoria, in which nuclear loci share large amounts of ancestral variation. In the case of nuclear genes of Lake Malawi, the absence of phylogenetically informative variation suggests a relative absence of ancestral variation. Nuclear genes also differed from the mitochondria in having nearly twice the amount of divergence from Oreochromis (tilapia). An approximate splitting time between mbuna and nonmbuna lineages was estimated as 0.7 Myr. Oreochromis is estimated to have diverged from the cichlids in Lake Malawi and Lake Tanganyika about 18 MYA.     

The endemic gastropod fauna of Lake Titicaca: correlation between molecular evolution and hydrographic history

Lake Titicaca, situated in the Altiplano high plateau, is the only ancient lake in South America. This 2- to 3-My-old (where My is million years) water body has had a complex history that included at least five major hydrological phases during the Pleistocene. It is generally assumed that these physical events helped shape the evolutionary history of the lake's biota. Herein, we study an endemic species assemblage in Lake Titicaca, composed of members of the microgastropod genus Heleobia, to determine whether the lake has functioned as a reservoir of relic species or the site of local diversification, to evaluate congruence of the regional paleohydrology and the evolutionary history of this assemblage, and to assess whether the geographic distributions of endemic lineages are hierarchical. Our phylogenetic analyses indicate that the Titicaca/Altiplano Heleobia fauna (together with few extralimital taxa) forms a species flock. A molecular clock analysis suggests that the most recent common ancestor (MRCAs) of the Altiplano taxa evolved 0.53 (0.28-0.80) My ago and the MRCAs of the Altiplano taxa and their extralimital sister group 0.92 (0.46-1.52) My ago. The endemic species of Lake Titicaca are younger than the lake itself, implying primarily intralacustrine speciation. Moreover, the timing of evolutionary branching events and the ages of two precursors of Lake Titicaca, lakes Cabana and Ballivián, is congruent. Although Lake Titicaca appears to have been the principal site of speciation for the regional Heleobia fauna, the contemporary spatial patterns of endemism have been masked by immigration and/or emigration events of local riverine taxa, which we attribute to the unstable hydrographic history of the Altiplano. Thus, a hierarchical distribution of endemism is not evident, but instead there is a single genetic break between two regional clades. We also discuss our findings in relation to studies of other regional biota and suggest that salinity tolerance was the most likely limiting factor in the evolution of Altiplano species flocks.     

Effects of the level of early productivity on the lifespan of ewes in contrasting flock environments

Selection for high levels of prolificacy has allowed substantial improvements in the production efficiency of New Zealand (NZ) sheep farms, but the consequences on ewe lifetime performance are mostly unknown. In this study, the relationship between the level of prolificacy early in ewes’ productive lives and their probability to survive later (i.e. stayability) was evaluated in two contrasting NZ flock environments. Records were obtained from 6605 ewes from four ram breeder flocks representing either a moderate (n=2) or a highly variable (n=2) nutritional environment. All ewes lambed for the first time at 2 years of age and were mated the following year. The number of lambs born during the first 2 years of productive life (NLB_2–3) was used as a measure of early prolificacy. Effects of NLB_2–3 on stayability to 4, 5, 6, 7 and 8 years old were analysed using logistic regression. Curvilinear effects (logit-transformed) were detected (P<0.05) until stayability to 6 years and to 8 years old in the highly variable and the moderate environment, respectively. The NLB_2–3 that resulted in maximum expected stayability to various ages was 3.9 to 4.2, and 4.5 to 4.7 lambs in the highly variable and in the moderate flock environments, respectively. In addition, ewe stayability was reduced when the proportion of the litter that survived from birth to weaning (i.e. ewe rearing ability) was submaximal during the early productive life. High prolific ewes had a low rearing ability whatever the environment whereas the rearing ability of lowly prolific ewes was apparently more sensitive to the nutritional environment. The poor maternal performance of ewes with low levels of NLB_2–3 led to a premature culling by breeders whereas the high early reproductive effort associated with high levels of NLB_2–3 seemed to be at the cost of ewes’ survival, even in the moderate flock environment. In conclusion, the flock environment influenced the level of early prolificacy beyond which ewe longevity was reduced. It is suggested that further selection for high and early prolificacy in NZ flocks is likely to impair ewes’ lifetime productivity.     

Kleptoparasitic prey competition in shoaling fish: effects of familiarity and prey distribution

Familiarity is thought to stabilize dominance hierarchies andreduce aggressive interactions within groups of socially livinganimals. Though familiarity has been widely studied in shoalingfish, few studies have investigated changes in prey competitionas a function of time spent together within groups of initiallyunfamiliar individuals. In this study, we created shoals ofthree-spined stickleback (Gasterosteus aculeatus) and monitoredchanges in foraging rates and related competitive behaviorswithin shoals over a 4-week period in experimental series whereprey was spatially and temporally concentrated or dispersed.Prey share was unequal under both prey distribution modes, anddisparity in prey share was not seen to change as trials progressed.Interestingly, the contest rate for prey items fell over timewhen individuals were competing for dispersed prey but not whenprey were concentrated. We found no evidence that fish showedassociation preferences for either group members that had consumeda greater or lesser proportion of prey during trials. Thoughthe intensity of competition may be reduced by increased groupstability in nature, this is likely to be strongly dependenton the way prey resources are distributed through space andtime.     

TESTS FOR ANCIENT SPECIES FLOCKS BASED ON MOLECULAR PHYLOGENETIC APPRAISALS OF SEBASTES ROCKFISHES AND OTHER MARINE FISHES

The concept of species flocks has been central to previous interpretations of patterns and processes of explosive species radiations within several groups of freshwater fishes. Here, molecular phytogenies of species-rich Sebastes rockfishes from the northeastern Pacific Ocean were used to test predictions of null theoretical models that assume random temporal placements of phylogenetic nodes. Similar appraisals were conducted using molecular data previously published for particular cichlid fishes in Africa that epitomize, by virtue of a rapid and recent radiation of species, the traditional concept of an intralacustrine “species flock.” As gauged by the magnitudes of genetic divergence in cytochrome b sequences from mitochondrial DNA, as well as in allozymes, most speciation events in the Sebastes complex were far more ancient than those in the cichlids. However, statistical tests of the nodal placements in the Sebastes phylogeny suggest that speciation events in the rockfishes were temporally nonrandom, with significant clustering of cladogenetic events in time. Similar conclusions also apply to an ancient complex of icefishes (within the Notothenioidei) analyzed in the same fashion. Thus, the rockfishes (and icefishes) may be interpreted as ancient species flocks in the marine realm. The analyses exemplified in this report introduce a conceptual and operational approach for extending the concept of species flocks to additional environmental settings and evolutionary timescales.