A review of Afrotropical nightjar mortality,mainly road kills

Afrotropical nightjars have evolved a number of adaptations that enable them to cope well with various natural mortality factors facing them. They are, however, extremely vulnerable to the many human factors that affect their lives. Three areas of human activity are of major concern: road traffic, habitat destruction and food gathering. Suitable legislation, with strict enforcement, should ensure that nightjar populations are not harvested excessively for the table. The many National Parks, Game Reserves and other protected areas throughout the Afrotropics should ensure that sufficient breeding and feeding habitats are available for the various nightjar species. There does not appear to be a solution to the road traffic problem, so road kills will almost certainly be the major mortality factor affecting most adult nightjars. A subjective assessment was made of the road mortality suffered by each species. The forest species (Brown Nightjar, Veles binotatus, Bates's Nightjar, Caprimulgus batesi and Prigogine's Nightjar, C. prigoginei) appear to be unaffected. The level of road mortality was rated as very low for Swamp Nightjar, C. natalensis, Nechisar Nightjar, C. solala, Golden Nightjar, C. eximius and Egyptian Nightjar, C. aetfvptius; as low for Montane Nightjar, C. poliocephalus, Slender-tailed Nightjar, C. clarus, Star-spotted Nightjar, C. stellatush and Red-necked Nightjar, C. ruficollis; as moderate for Fiery-necked (Pectoral) Nightjar, C. pectoralis, Donaldson-Smith's Nightjar, C. donaldsoni, Dark Nightjar, C. fraenatus and Nubian Nightjar, C. nubicus; as high for Freckled Nightjar, C. tristigma, Rufous-cheeked Nightjar, C. rufigena, Plain Nightjar, C. inornatus and Standard-winged Nightjar, Macrodipteryx longipennis; and as very high for Long-tailed Nightjar, C. climacurus, Square-tailed Nightjar, C. fossii, European Nightjar, C. europaeus and Pennant-winged Nightjar, Semeiophorus vexillarius. Throughout the Afrotropical Region nightjars die on the roads in great numbers and for some species the road toll is a major mortality factor.     

Africa Climate exchange

Golden Nightjar Caprimulgus eximius is an apparently sedentary sub-Saharan species with a breeding range extending from Senegal and Mauritania to Sudan. Although genetic studies of nightjars and related Caprimulgiformes have been published previously, none has included Golden Nightjar. In this study, mitochondrial and nuclear DNA of a Golden Nightjar found dead in Western Sahara in April 2016 was sequenced and compared with other species in the genus Caprimulgus. It was concluded with strong support that Golden Nightjar is closely related to Egyptian Nightjar C. aegyptius. It is hypothesised that Golden and Egyptian Nightjars may have arisen by splitting of a single ancestral species into migratory and sedentary populations.     

Environmental correlates of Freckled Nightjar ( Caprimulgus tristigma ) activity in a seasonal, subtropical habitat

We examined environmental correlates of activity in the Freckled Nightjar (Caprimulgus tristigma), a nocturnal aerial insectivore that is resident year-round in the colder and drier parts of southern Africa. Specifically, we tested the prediction that air temperature (T _a), in addition to light availability, is a significant correlate of Freckled Nightjar activity. We found that aerial insect density varied by over three orders of magnitude (from <1.0 to 117.1 insects 1,000 m⁻³) during the course of the study, and was strongly and positively related to T _a. Nightjar activity was also significantly temperature-dependent, with a cessation of activity at T _a < 12°C. Consistent with previous studies, we found that activity levels were strongly related to ambient light, with markedly reduced activity levels on dark nights. The average number of Freckled Nightjars encountered per 14.6-km transect was 1.95 ± 2.44 nightjars transect⁻¹ during nights near full moon, but only 0.17 ± 0.61 nightjars transect⁻¹ around new moon. Our study reveals that caprimulgid activity can be significantly influenced by temperature as well as ambient light.     

Comparison of the two rupicolous nightjars: the Afrotropical Freckled Nightjar Caprimulgus tristigma and the Neotropical Blackish Nightjar C. nigrescens

The Freckled Nightjar Caprimulgus tristigma and the Blackish Nightjar C. nigrescens are widespread and common within their rupicolous habitat, in the Afrotropics and Neotropics respectively, and may therefore be considered as successful in their adaptation to this habitat, a niche that has not been exploited by any other nightjar species. However, apart from a plumage pattern that matches a rocky substrate, their known life histories provide no common factors to explain this adaptive success. The factors that they do share are common to most other nightjars. While they nest and roost on rocks, their breeding biology is remarkably different. The contrasts and lack of convergence are surprising, and suggest that these two species are not as closely related as their current congeneric status implies. This is supported by recent molecular studies that place the African and South American Caprimulgus species in different well-supported clades.     

DNA diet profiles with high‐resolution animal tracking data reveal levels of prey selection relative to habitat choice in a crepuscular insectivorous bird

Given the global decline of many invertebrate food resources, it is fundamental to understand the dietary requirements of insectivores. We give new insights into the functional relationship between the spatial habitat use, food availability, and diet of a crepuscular aerial insectivore, the European Nightjar (Caprimulgus europaeus) by relating spatial use data with high‐throughput sequencing (HTS) combined with DNA metabarcoding. Our study supports the predictions that nightjars collect a substantial part of their daily nourishment from foraging locations, sometimes at considerable distance from nesting sites. Lepidopterans comprise 65% of nightjars'' food source. Nightjars tend to select larger species of Lepidoptera (>19 mm) which suggests that nightjars optimize the efficiency of foraging trips by selecting the most energetically favorable—larger—prey items. We anticipate that our findings may shed additional light on the interactions between invertebrate communities and higher trophic levels, which is required to understand the repercussions of changing food resources on individual‐ and population‐level processes.     

Study on the foraging behaviour of the European nightjar Caprimulgus europaeus reveals the need for a change in conservation strategy in Belgium

Effective nature conservation requires coherent actions based on the best available evidence concerning protected species. Recent studies have suggested that European nightjars Caprimulgus europaeus forage outside their recognized breeding habitats, yet, for Flanders (northern Belgium) information on nightjar foraging behaviour and key foraging habitats is lacking. To assess whether the foraging ecology of nightjars in Flanders is similar to that observed in other parts of Europe, we studied the crepuscular behaviour of this species in Bosland (northeastern Flanders) during a five‐year radio telemetry study. Tracking of 48 individuals within a coniferous forest was standardized and home ranges were calculated using a kernel density estimator (fixed kernel). Habitat use was investigated by comparing kernel placement to available habitat. Average maximal foraging distance was 2603 ± 1094 m and home ranges extended up to 691 ha. We identified the key foraging habitats to be extensively‐cultivated grasslands and recreational areas, areas that were previously assumed unsuitable for Belgian nightjars. Our results indicate the importance of foraging sites outside the breeding territory, confirming the findings of previous studies performed elsewhere in Europe. Incorporating our findings into future conservation plans could, therefore, lead to improved efficiency of EU conservation measures, designed for the protection of this bird species in Flanders.     

ASPECTS OF THE BREEDING BIOLOGY OF THE FIERYNECKED NIGHTJAR

Jackson H. D. 1985. Aspects of the breeding biology of the Fierynecked Nightjar. Ostrich 56: 263–276.

A marked population of nightjars in Zimbabwe was studied intensively for four breeding seasons. This paper covers certain aspects of the breeding biology of the Fierynecked Nightjar Caprimulgus pectoralis. The male shows strong site fidelity during the breeding season (September to December), singing, feeding and breeding within an area of about 5,8 ha. There is some evidence of site defence by the male. The female shows strong mate fidelity, resulting in a pair bond for life. Egg laying starts with full moon in September and is further stimulated by the next two full moon periods. The eggs are laid directly on dense leaf litter at a site overhung by foliage. The normal clutch is two eggs (12S % are one egg) laid on successive days during the afternoon. Incubation starts with the first egg and is by the male at night and the female by day. The incubation period is 18 days. The birds respond to undue disturbance by deserting the eggs and laying a replacement clutch. The chicks usually hatch on successive afternoons; they are mobile on the first day and will move to a parent if called. Both parents feed and brood the young during twilight and moonlight; the male broods them on dark nights and the female does so by day. The species is double-brooded when time permits, the female laying again once the first brood has reached independence; she may even lay a third clutch if the second one comes to grief. There is no evidence of adults transporting eggs or young.     

The Use of Automated Bioacoustic Recorders to Replace Human Wildlife Surveys: An Example Using Nightjars

To be able to monitor and protect endangered species, we need accurate information on their numbers and where they live. Survey methods using automated bioacoustic recorders offer significant promise, especially for species whose behaviour or ecology reduces their detectability during traditional surveys, such as the European nightjar. In this study we examined the utility of automated bioacoustic recorders and the associated classification software as a way to survey for wildlife, using the nightjar as an example. We compared traditional human surveys with results obtained from bioacoustic recorders. When we compared these two methods using the recordings made at the same time as the human surveys, we found that recorders were better at detecting nightjars. However, in practice fieldworkers are likely to deploy recorders for extended periods to make best use of them. Our comparison of this practical approach with human surveys revealed that recorders were significantly better at detecting nightjars than human surveyors: recorders detected nightjars during 19 of 22 survey periods, while surveyors detected nightjars on only six of these occasions. In addition, there was no correlation between the amount of vocalisation captured by the acoustic recorders and the abundance of nightjars as recorded by human surveyors. The data obtained from the recorders revealed that nightjars were most active just before dawn and just after dusk, and least active during the middle of the night. As a result, we found that recording at both dusk and dawn or only at dawn would give reasonably high levels of detection while significantly reducing recording time, preserving battery life. Our analyses suggest that automated bioacoustic recorders could increase the detection of other species, particularly those that are known to be difficult to detect using traditional survey methods. The accuracy of detection is especially important when the data are used to inform conservation.     

Effects of light and prey availability on nocturnal, lunar and seasonal activity of tropical nightjars

Nightjars and their allies represent the only major group of visually hunting aerial insectivores with a crepuscular and/or nocturnal lifestyle. Our purpose was to examine how both light regime and prey abundance in the tropics, where periods of twilight are extremely short, but nightjar diversity is high, affect activity across different temporal scales. We studied two nightjar species in West African bush savannah, standard‐winged nightjars Macrodipteryx longipennis Shaw and long‐tailed nightjars Caprimulgus climacurus Vieillot. We measured biomass of potential prey available using a vehicle mounted trap and found that it was highest at dusk and significantly lower at dawn and during the night. Based on direct observations, both nightjars exhibit the most intense foraging behaviour at dusk, less intense foraging at dawn and least at night, as predicted by both prey abundance and conditions for visual prey detection. Nocturnal foraging was positively correlated with lunar light levels and ceased below about 0.03 mW m^?2. Over the course of a lunar cycle, nocturnal light availability varied markedly, while prey abundance remained constant at dusk and at night was slightly higher at full moon. Both species increased twilight foraging activity during new moon periods, compensating for the shorter nocturnal foraging window at that time. Seasonally, the pattern of nocturnal light availability was similar throughout the year, while prey availability peaked shortly after onset of the wet season and then slowly decreased over the following four months. The courtship and breeding phenology of both species was timed to coincide with the peak in aerial insect abundance, suggesting that prey availability rather than direct abiotic factors act as constraints, at least at the seasonal level. Our findings illustrate the peculiar constraints on visually orienting aerial nocturnal insectivores in general and tropical nightjars in particular and highlight the resulting nocturnal, lunar and seasonal allocation of activities.     

Human-Induced Changes in Landscape Configuration Influence Individual Movement Routines: Lessons from a Versatile,Highly Mobile Species

Landscape conversion by humans may have detrimental effects on animal populations inhabiting managed ecosystems, but human-altered areas may also provide suitable environments for tolerant species. We investigated the spatial ecology of a highly mobile nocturnal avian species–the red-necked nightjar (Caprimulgus ruficollis)–in two contrastingly managed areas in Southwestern Spain to provide management recommendations for species having multiple habitat requirements. Based on habitat use by radiotagged nightjars, we created maps of functional heterogeneity in both areas so that the movements of breeding individuals could be modeled using least-cost path analyses. In both the natural and the managed area, nightjars used remnants of native shrublands as nesting sites, while pinewood patches (either newly planted or natural mature) and roads were selected as roosting and foraging habitats, respectively. Although the fraction of functional habitat was held relatively constant (60.9% vs. 74.1% in the natural and the managed area, respectively), landscape configuration changed noticeably. As a result, least-cost routes (summed linear distances) from nest locations to the nearest roost and foraging sites were three times larger in the natural than in the managed area (mean ± SE: 1356±76 m vs. 439±32 m). It seems likely that the increased proximity of functional habitats in the managed area relative to the natural one is underlying the significantly higher abundances of nightjars observed therein, where breeders should travel shorter distances to link together essential resources, thus likely reducing their energy expenditure and mortality risks. Our results suggest that landscape configuration, but not habitat availability, is responsible for the observed differences between the natural and the managed area in the abundance and movements of breeding nightjars, although no effect on body condition was detected. Agricultural landscapes could be moderately managed to preserve small native remnants and to favor the juxtaposition of functional habitats to benefit those farm species relying on patchy resources.     

Diurnal body temperature patterns in free‐ranging populations of two southern African arid‐zone nightjars

Endotherms allocate large amounts of energy and water to the regulation of a precise body temperature (T_b), but can potentially reduce thermoregulatory costs by allowing T_b to deviate from normothermic levels. Many data on heterothermy at low air temperatures (T_a) exist for caprimulgids, whereas data on thermoregulation at high T_a are largely absent, despite members of this taxon frequently roosting and nesting in sites exposed to high operative temperatures. We investigated thermoregulation in free‐ranging rufous‐cheeked nightjars Caprimulgus rufigena and freckled nightjars Caprimulgus tristigma in the southern African arid zone. Individuals of both species showed labile T_b fluctuating around a single modal T_b (T_b‐mod). Average T_b‐mod was 39.7°C for rufous‐cheeked nightjars and 39.0°C for freckled nightjars. In both species, diurnal T_b increased with increasing T_a. At T_a ≥ 38°C, rufous‐cheeked nightjar mean T_b increased to 42°C, equivalent to 2.3°C above T_b‐mod. Under similar conditions, freckled nightjar T_b was on average only 1.1°C above T_b‐mod, with a mean T_b of 40.0°C. Freckled nightjars are one of the most heterothermic caprimulgids investigated to date, but our data suggest that during hot conditions this species maintains T_b within a narrow range above T_b‐mod, possibly reflecting an evolutionary tradeoff between decreased thermal sensitivity to lower T_b but increased sensitivity to high T_b. These findings reveal how general thermoregulatory patterns at similar T_a can vary even among closely related species.     

THE ENGLISH NAMES OF THE AFROTROPICAL NIGHTJARS (CAPRIMULGIDAE)

Jackson, H. D. 1993. The English names of the Afrotropical nightjars (Caprimulgidae). Ostrich 64: 148–159.

At least 172 different English names have been used on the nightjars of Africa and its related islands, including Madagascar. A review of the literature chronicles the name changes within each species. Problems created by some duplication and transposition of names between species are discussed. A key to all the names is provided by an alphabetical list cross referenced to the species concerned. The orthography of names is discussed and it is recommended that all animal names be capitalized and freed of unnecessary hyphens. The nightjar names used in The Birds of Africa are accepted as the standard to be followed subject to a few changes.     

Vocal individuality of large-tailed nightjar (Caprimulgus macrurus) in Peninsular Malaysia

All four nightjar species resident in Peninsular Malaysia are vocally distinct, yet they remain little studied. Conventional field methods based on visual cues to study diurnal species may be impractical for nightjars. Alternatively, aural survey can potentially be applied on nightjars provided that individuality in their vocalisations can be proven. Our study aimed to determine the vocal individuality of the common, large-tailed nightjar (Caprimulgus macrurus) in oil palm smallholdings and an isolated forest patch located in Selangor, Peninsular Malaysia. From the call recordings obtained from 22 individuals, results of the Kruskal-Wallis analysis revealed significant differences in all the nine vocal parameters (call length, interquartile range bandwidth, low, high, average, centre and peak frequencies as well as first and third quartile frequencies) measured among individual nightjars (p < 0.001) regardless of study sites. Discriminant Function Analysis showed that more than 94.5% of original grouped cases were correctly classified. This implied that the majority of vocalizations can be assigned to individual birds based on the parameters measured. This study demonstrated the occurrence of vocal individuality in the large-tailed nightjar and such a finding pertaining to distinct vocalisations at the individual level will compensate for the limited access to visual cues in field surveys, as with the case for all nocturnal birds.     

Home‐range size and habitat use of European Nightjars Caprimulgus europaeus nesting in a complex plantation‐forest landscape

In Europe, the consequences of commercial plantation management for birds of conservation concern are poorly understood. The European Nightjar Caprimulgus europaeus is a species of conservation concern across Europe due to population depletion through habitat loss. Pine plantation‐forest is now a key Nightjar nesting habitat, particularly in northwestern Europe, and increased understanding of foraging habitat selection is required. We radiotracked 31 Nightjars in an extensive (185‐km²) complex conifer plantation landscape in 2009 and 2010. Home‐range 95% kernels for females, paired males and unpaired males were an order of magnitude larger than song territories of paired males, emphasizing the importance of habitats beyond the song territory. Nightjars travelled a mean maximum distance of 747 m from the territory centre each night. Home‐range placement relative to landscape composition was examined by compositional analysis. Pre‐closure canopy forest (aged 5–10 years) was selected at all scales (MCP, 95% and 50% kernels), with newly planted forest (aged 0–4 years) also selected within 50% kernels. For telemetry fixes relative to habitat composition within 2 km of their territory centre, individuals again selected pre‐closure and newly planted forest, and also grazed grass heath. Open ungrazed habitat was not selected, with implications for open habitat planning for biodiversity conservation within public‐owned forests. Despite the Nightjars’ selection for younger growth, moth biomass was greater in older forest stands, suggesting that foraging site selection reflects ease of prey capture rather than prey abundance. Within large plantation‐forest landscapes, a variety of growth stages is important for this species and our results suggest that grazing of open habitats within and adjacent to forest will additionally benefit the European Nightjar.     

Observation of nest visitation by multiple males in the European Nightjar Caprimulgus europeaus

ABSTRACT

We report observations of nest visitation of multiple males to three nests of European Nightjars Caprimulgus europeaus. In one case, we report the direct observation of two males apparently exhibiting parental behaviour at the same nest. In two other cases, second males visited nests. We consider the potential functions of nest visits by extra-pair males in the Nightjar.     

Genomic investigation refutes record of most diverged avian hybrid

The most diverged avian hybrid that has been documented (Numida meleagris × Penelope superciliaris) was reported in 1957. This identification has yet to be confirmed, and like most contemporary studies of hybridization, the identification was based on phenotype, which can be misleading. In this study, we sequenced the specimen in question and performed analyses to validate the specimen's parentage. We extracted DNA from the specimen in a dedicated ancient DNA facility and performed whole-genome short-read sequencing. We used BLAST to find Galliformes sequences similar to the hybrid specimen reads. We found that the proportion of BLAST hits mapped overwhelmingly to two species, N. meleagris and Gallus gallus. Additionally, we constructed phylogenies using avian orthologs and parsed the species placed as sister to the hybrid. Again, the hybrid specimen was placed as a sister to N. meleagris and G. gallus. Despite not being a hybrid between N. meleagris and P. superciliaris, the hybrid still represents the most diverged avian hybrid confirmed with genetic data. In addition to correcting the “record” of the most diverged avian hybrid, these findings support recent assertions that morphological and behavioral-based identifications of avian hybrids can be error-prone. Consequently, this study serves as a cautionary tale to researchers of hybridization.     

Sequence variations in the large-subunit ribosomal RNA gene of ammonia (foraminifera,protozoa) and their evolutionary implications

An unusually high divergence was observed in the ribosomal RNA genes of a free-living population of foraminifera belonging to the genusAmmonia. The sequences of a large-subunit (LSU) rDNA expansion segment D1 and flanking regions were obtained from 20 specimens namedAmmonia sp. 1 andAmmonia sp. 2. The sequence divergence between the two species averages 14%. Within each species it ranges from 0.2% to 7.1% inAmmonia sp. 1 and from 0.7% to 2.3% inAmmonia sp. 2. We did not find two specimens having identical sequences. Moreover, in opposition to the generally acaepted view, rDNA sequence variations were also found within a single individual. The variations among several rDNA copies in a single specimen ofAmmonia may reach up to 4.9%. Most of the observed variations result from multiplication of CA or TA serial repeats occurring in two particularly variable regions. For single base changes, C-T transitions are most frequently observed. We discuss the evolution of expansion segments and their use for phylogenetic studies. Correspondence to: J. Pawlowski     

A field survey to investigate why nightjars frequent roads at night

Nightjars throughout the Afrotropical Region regularly frequent roads at night and during the twilight of dusk and dawn. Most species exhibit this behaviour and numerous individuals are killed by road traffic. Many theories have been advanced to explain the presence of nightjars on roads at night, but very little fieldwork has been carried out to test them. For a period of a year we carried out a weekly survey of nightjars on a selected road network near Harare in Zimbabwe. All nightjar encounters were documented in relation to variations in road surface, road width, adjacent habitat, arc of sky visible and other variables, such as the time of night, moon phase and weather conditions. None of these factors provided a complete explanation for the presence of nightjars on the roads surveyed. Neither were any of the nightjars seen dust bathing, taking grit or picking up insects from the road. Many were seen actively hunting flying insects, especially at dusk, and palpation of the stomachs of those caught confirmed that they had fed well during the evening hours. They were clearly using the road as an observation platform for hunting during the first few hours after sunset and then as a convenient place for resting and digesting. A nightjar sitting on an open road is certainly in a good position to see flying insects silhouetted against the twilight sky.     

Identification of different song types in the European Nightjar Caprimulgus europaeus

ABSTRACT

Capsule: Two distinct song types were identified for male European Nightjars Caprimulgus europaeus with their relative frequency of use changing through the breeding season, indicating a possible link to paired status.

Aims: To test whether two song types could be defined in audio recordings and whether use differed in relation to the paired status of males.

Methods: Unattended acoustic recording devices were placed at a Nightjar study site in Nottinghamshire, United Kingdom, and recordings of churring vocalizations were made during two periods of the breeding season. These recordings were then analyzed to identify the presence/absence of the song terminal phrase and associated audible features.

Results: Two distinct song types were identified in the recorded audio data that differed in their terminal phrasing and overall song duration. The number of Nightjar songs with a terminal phrase increased significantly between the two sampling periods, from lower levels during the site arrival period, to higher levels during the first clutch initiation period.

Conclusion: This study showed that the use of Nightjar song types appears to vary through the breeding season, with males being more likely to produce song with a terminal phrase during the first clutch initiation period, when they are more likely to be paired or in the presence of a female. The unattended acoustic recording method may provide a minimally intrusive means of assessing the number of Nightjar breeding pairs and not just singing males.     

Migration strategies and annual space‐use in an Afro‐Palaearctic aerial insectivore – the European nightjar Caprimulgus europaeus

Obligate insectivorous birds breeding in high latitudes travel thousands of kilometres during annual movements to track the local seasonal peaks of food abundance in a continuously fluctuating resource landscape. Avian migrants use an array of strategies when conducting these movements depending on e.g. morphology, life history traits and environmental factors encountered en route. Here we used geolocators to derive data on the annual space‐use, temporal pattern and migratory strategies in an Afro‐Palaearctic aerial insectivorous bird species – the European nightjar Caprimulgus europaeus. More specifically, we aimed to test a set of hypothesises pertaining to the migration of a population of nightjars breeding in south‐eastern Sweden. We found that the birds wintered across the central and western parts of the southern tropical Africa almost entirely outside the currently described wintering range of the species. The nightjars performed a narrow loop migration across Sahara, with spring Sahel stopovers significantly to the west of autumn stops indicative to an adaptive response to winds during migration. To our surprise, the migration speed was faster in the autumn (119 km d^{? 1}) than in the spring (99 km d^{? 1}), possibly due to the prevailing wind regimes over the Sahara. The estimated flight fraction in both autumn (14%) and spring (12%) was almost exactly as the theoretically predicted 1:7 time relationship between flights and stopovers for small birds. The temporal patterns within the annual cycle indicate that individuals follow alternative spatiotemporal schedules that converge towards the breeding season. The positive relationship between the spatially and temporally distant winter departure and breeding arrival suggests that individuals´ temporal fine‐tuning to breeding may be constrained, leading to potential negative fitness consequences.