Short-clawed Lark Certhilauda chuana Scientific name definitions

G. Derek Engelbrecht
Version: 2.0 — Published February 23, 2023

Breeding

Introduction

Short-clawed Lark is monogamous, the pair-bond typically lasting a single breeding season, followed by females dispersing during the non-breeding season. The breeding season starts in spring (August/September), peaks at the height of the wet season (December), and is generally completed by autumn (April). The nest is an open cup, constructed on the ground, at the base of vegetation, and is usually extremely well-concealed. The clutch size is 2–3 eggs, but there is seasonal clutch size variation; larger clutches are produced during and shortly after the peak of the wet season. The incubation period is 15 d and is carried out by the female only, with the male acting as a sentinel to warn of imminent danger near the nest. The altricial nestlings are fed invertebrates by both parents and leave the nest after 11–12 d. Little is known about the behavior of fledglings and the duration of the post-fledging period.

Phenology

In Botswana, egg-laying spans four months, from September–January, with a peak in October and November (30, 6, 31). This period coincides with the early part of the wet season, with breeding largely completed before the peak of the wet season (December–February) (6). On the Polokwane Plateau, egg-laying spans mid-August–mid-March, with a peak from October–December (5, 46); breeding in this population is more evenly spaced over the seven-month breeding season, but egg-laying peaks at or just after the peak of the wet season (December) (5). The differences observed in the breeding seasons between the two regions may, however, be artificial, as the population on the Polokwane Plateau has been monitored more intensely. In both populations, analysis of the breeding phenology shows a bimodal pattern, with an early-season and main-season peak in the initiation of nesting attempts. The low number of nests initiated between the two peaks reflects pairs still actively nesting or attending to fledged young from the early-season peak, with most of the new nests initiated in this period being replacement clutches or perhaps late starters (6, 5; DE, unpublished data). There is, however, a reasonable amount of inter-annual variation in the onset of laying in both populations, with the earliest recorded onset of breeding on the Polokwane Plateau initiated 54 d after the northern solstice (i.e., 14 August), and the latest onset of breeding 129 d after the northern solstice (i.e., 28 October) (46). It has been suggested that the onset of nesting may be linked to local rainfall events (6), but the mid-August record on the Polokwane Plateau occurred weeks before the first rain in that season (DE, unpublished data).

Nest Site

Selection Process

Nest-site selection is presumably by the female as nest construction is solely performed by the female.

Microhabitat

The nests are almost invariably placed at the base of a tuft of grass (e.g., Cymbopogon sp., Eragrostis curvula, Themeda triandra, Cynodon dactylon, Digitaria milinjiana), a small forb (e.g., Jatropha zeyheri, Geigeria burkei, Chamaecrista mimosoides, Hypoxis rigidula, Chrysocoma tenuifolia, Melhania sp.), or a small bush (Vachellia tortilis, Vachellia [Senegalia] mellifera) (47, 6, 5). A minority (3.5% on the Polokwane Plateau) of nests are completely exposed (5). In Botswana, seven of 12 nests (58%) were placed below a bush or forb, with the remainder placed at the base of or between two adjacent grass tufts; on the Polokwane Plateau, most nests were placed at the base of a grass tuft (79%), and relatively few were associated with a forb or woody shrub (6; DE, unpublished data). These differences may be due to the different habitats in which the studies were conducted: fallow fields or heavily-grazed areas with pioneer plants in rural subsistence farmland in Botswana versus the relatively pristine Polokwane Game Reserve on the Polokwane Plateau.

Site Characteristics

Although territories are characterized by short grass with a lot of bare ground (6, 41), nest microhabitat selection is usually in areas with a fair amount of basal cover (~40%) within a territory (DE, unpublished data). Although nest entrances were recorded facing all directions, nest entrances are not randomly distributed. There is a tendency for nests to face north in Botswana (6) or north-northwest on the Polokwane Plateau (DE, unpublished data), but the proximate reasons for this tendency is not known. A majority of nests are well-concealed by the base plant that, in addition to concealment, also provides shade to the eggs, nestlings, or attending parent for at least part of the day. However, this is not a prerequisite, as some nests are only moderately concealed or wholly exposed. The mainly northerly orientation of nest entrances suggests shading of the nest is perhaps not a critical factor in nest site and entrance selection. Nests are usually placed in the vicinity (5–15 m) of a small tree or bush used by males to perform sentinel duties while the female incubates (DE, unpublished data).

Nest

Construction Process

Females are solely responsible for nest construction and maintenance (5). During nest construction, the male follows the female closely and will act as a sentry by perching and vocalizing from a nearby perch while the female is collecting nest material (5). Nest construction, defined as the period from the start of nest construction to laying the first egg, lasts 4–5 d but can be as long as 14 d (5). Construction is mainly confined to the mornings. The female will continue to add lining throughout the incubation period and occasionally in the early stages of the nestling period (DE, unpublished data). Nest material is often carried away or consumed by termites, sometimes presenting an easy meal for the parents (6). On one occasion, termites constructed a soil sheet over the nest material on the cup lip, and the entire nest structure, except for the lining, was eventually covered in the soil sheet.

Structure and Composition

The nest is a shallow, open cup in the ground. The nest structure consists mainly of coarse grass leaf blades, grass culms, and grassroots, while the nest lining consists of more delicate grass blades, rootlets, and grass panicles. Other materials recorded in nests include small numbers of contour feathers, the leaves of herbaceous plants, twigs, bits of bark, thorns of Vachellia spp., hair, and pieces of livestock dung (5, 6; DE, unpublished data). There is considerable variation in the structure of nests, with some containing barely any lining and others having a well-developed lining and an apron (5). Approximately 50% of nests on the Polokwane Plateau have a well-developed apron of grass culms, panicles, stems, and roots at the entrance (DE, unpublished data). Some nests have a slightly raised lip, which may be due to the accumulation of material during the excavation of the cup. In at least one instance, it prevented the nest from getting flooded during a heavy storm (DE, unpublished data).

Dimensions

The mean nest dimensions are: cup diameter, 98 mm (range: 53–90 mm), cup depth, 35 mm (range: 20–60 mm) (6, 5; DE, unpublished data). The mass of nests varies considerably (mean of 25.7 g, range 3.5–64.0 g) (5; DE, unpublished data).

Maintenance or Reuse of Nests

A new nest is constructed for each new nesting attempt, sometimes as close as 5 m from an earlier nest within the same season (DE, unpublished data).

Eggs

Shape

The eggs are oval to slightly elliptical (DE, unpublished data).

Size

Mean dimensions (length x width): 21.87 mm ± 0.88 SD (range 19.79–24.55) × 15.65 mm ± 0.44 SD (range 14.50–16.72; n = 85 eggs) (DE, unpublished data). There is no difference in egg sizes between the western and eastern populations (6, 5).

Mean egg Elongation Index (l/w): 1.40 ± 0.05 SD (range 1.28–1.54; n=85) (DE, unpublished data).

Mean egg volume (according to 51): 2,734.03 mm3 ± 217.05 SD (range 2,169.12–3,216.49; n = 85) (DE, unpublished data).

Mass

The mean fresh egg mass is 2.74 g ± 0.22 SD (range 2.10–3.23, n = 37) (DE, unpublished data). The mean fresh egg mass represents 8.8% (range 6.9–10.6, n = 24) of the mean mass of adult females.

Eggshell Thickness

Information needed.

Color and Surface Texture

The base color of eggs is off-white to buff, rarely buff-brown, and they are liberally spotted with shades of brown, sepia, and gray. Most eggs have spots concentrated at the larger end, with the larger end of some eggs being almost uniformly brown. The surface of the egg is smooth and matte.

Clutch Size

Clutch size ranges from 1–3 eggs, but clutch sizes vary seasonally and possibly regionally. The mean clutch size in Botswana is 2.75 eggs (n = 12 nests; 6), and on the Polokwane Plateau it is 2.22 ± 0.42 SD (n = 54 nests) (DE, unpublished data), suggesting regional clutch-size variation may exist (5). Although based on a small sample of only 12 nests, the brevity of the breeding season in Botswana (four months) suggests populations in that region may not exhibit seasonal variation in clutch size and they may also show fewer nesting attempts but larger clutch sizes. In contrast, the eastern population on the Polokwane Plateau is multi-brooded, lays smaller clutches, and shows seasonal clutch-size variation. On the Polokwane Plateau, mean clutch sizes are smallest (mean of 2.0 eggs) at the beginning and end of the breeding season (5). There is a steady increase in clutch sizes from the start of the wet season in October/November, reaching a peak in January (mean of 2.67 eggs) and February (mean of 2.5 eggs), just after the peak of the wet season in this region (DE, unpublished data); although fewer new breeding attempts are initiated in January and February, the mean clutch size is larger during this period (DE, unpublished data). Two single-egg clutches recorded on the Polokwane Plateau were at the beginning of the breeding season, and both were replacement clutches following earlier nest failure.

Egg Laying

Eggs are laid in the mornings, usually before 0800 h, at one-day intervals (5). During laying, the female will occasionally visit the nest to deliver lining. After nest failure or fledging of a brood, the female may start constructing a new nest within two days and start laying again within a week (5). It is not uncommon to see an incubating female still feeding fledglings from a previous brood (DE, unpublished data).

Incubation

Onset of Broodiness and Incubation in Relation to Laying

Full-contact incubation starts upon clutch completion (5). Females deliver lining during egg-laying and may perform non-contact incubation for short spells (DE, unpublished data).

Incubation Period

The mean incubation period lasts 15.1 d ± 0.99 SD (range 14–16, n = 8) (DE, unpublished data).

Parental Behavior

Incubation is by the female alone, while the male undertakes sentinel duties near the nest, calling and frequently singing (5). In the face of a potential threat, males utter contact or alarm calls, and females slip off nests and fly away or run through the grass and take off a few meters from the nest (DE, unpublished data).

Nests are attended on average 70.9% of the time during incubation (n = 8 nests) (DE, unpublished data). Incubation bouts are relatively short in the early- to mid-morning but get longer as the day progresses. From mid-morning to mid-afternoon, nest attendance is near-constant, with the female shading the eggs for extended periods. In sweltering conditions, females are reluctant to leave the nest in the face of disturbance (6). The average duration of a complete incubation on-bout, i.e., one where the incubating bird was not disturbed by the observer or equipment failure resulted in an incomplete estimate of the duration, is 41.33 min ± 62.62 SD (median 25.5 min, range 5–341; n = 30). The longest recorded bout lasted at least 469 min before equipment failure (DE, unpublished data). The mean duration of an incubation recess bout is 21.25 min ± 16.93 SD (median 16.0 min, range 1–64; n = 29) (DE, unpublished data).

During incubation, the female would turn the eggs and adjust her position fairly regularly. In hot conditions, the female would employ various ways to dissipate body heat, e.g., panting, holding the wings slightly away from the body, raising the crown, back, and rump feathers, and closing the eyes to slits, presumably to reduce water loss from the surface of the eye. Under such conditions, she would also stand above the eggs to shade them rather than perform contact incubation (DE, unpublished data).

No provisioning of the incubating female by the male has been recorded.

Hardiness of Eggs Against Temperature Stress

Information needed.

Hatching

Shell Breaking and Emergence

Hatching is synchronous on the same day, but the individual nestlings may hatch over several hours. Nestlings usually hatch within two hours of each other, but sometimes as long as eight hours may elapse between the first and last hatched nestlings (5). The hatching period, from pipping to emerging from the eggs, is 2–4 h (n = 1) (5).

Parental Assistance and Disposal of Eggshells

The female disposes of eggshells soon after hatching.

Young Birds

Condition at Hatching

Newly hatched (day zero) nestlings have tufts of long, pale, straw-colored or buff down on all the dorsal pterylae, but the underparts are naked. The dorsal skin surface is a dark purplish color and the ventral surface is a paler purplish-pink. The culmen is horn-colored and the gape flanges are pale creamy-yellow, almost white. The inside of the mouth is bright orange with two lateral, oval to elliptical, black tongue spots towards the posterior part of the tongue. There is no apical tongue spot, but there are two spots at the tips of the mandible and maxilla. The eyes are closed and none of the feather tracts are visible below the skin. The mean mass of nestlings measured on the hatching day is 2.52 g ± 0.14 SD (range 2.36–2.60, n = 3).

Growth and Development

The nestling period is 11–12 d (6, 5). On day one, the feather tracts are visible below the skin, but no feathers have erupted through the skin. On day two, the first feathers on the spinal, femoral, and ventral tracts have erupted through the skin, and by day three, all the feather tracts have at least some feathers in pin. The eyes begin to open and appear as slits on day three. On day four, the first feathers on the capital, spinal, ventral, femoral, and crural tracts are in brush, and, although the eyes can open reasonably wide, they are mostly kept closed. By day five, the eyes are fully open, and the nestlings make an audible but high-pitched tseep begging call. The first secondaries, secondary coverts, and rectrices are in brush on day six, but the first primaries are only in brush a day later (day seven). Days eight and nine see more of the primaries, primary-coverts, secondaries, and secondary coverts in brush. By the end of day nine, all the inner secondaries and secondaries are in brush, and about two-thirds of the primaries. By day 10, all the primaries are in brush and most of the primary coverts. At around fledging (~day 11), all the primaries are between 30% and 66% in brush (5). The fledglings retain some down above the supercilium and towards the femoral region for a few days after fledging.

When nestlings are left unattended, they lie silent and motionless with their eyes closed or open only as slits, typically only moving, gaping, and begging when parents call the contact feeding call. On one occasion, video footage showed a stray dog discovering a nest with two 10-day-old nestlings, and despite the dog sniffing at the nestlings, they remained motionless . It appeared to have been an effective strategy as the dog left without attempting to depredate the nestlings. During the latter stages of the nestling period, heat-stressed nestlings may pant for extensive periods and position themselves to maximize the amount of shade they are exposed to.

Parental Care

Short-clawed Lark exhibits biparental care, but there exist clear role divisions. Females are solely responsible for nest construction, incubation, and nest maintenance, while the males act as sentinels near the nest to warn of potential threats. The pair share provisioning, nest sanitation and, to a lesser extent, brooding duties. However, the individual contributions of males are quite variable, with some males matching or exceeding the provisioning rate of their mates, while others only occasionally deliver food (6, 5). Although brooding by males was not reported by Herremans and Herremans (6) or Engelbrecht (5), a study using video footage revealed some males occasionally assist with brooding (DE, unpublished data).

Adults are wary near the nest. When returning to the nest to incubate, brood, or provision the young, they typically perch nearby, scan the surroundings for several minutes, and give the contact call (5). If all appears safe, the returning parent will land a few meters from the nest and course their way towards the nest. Upon arrival at the nest, they give the agitated contact call to stimulate the nestlings to gape and feed them. When leaving, they occasionally take flight from just outside the nest, but more typically walk at least a meter, sometimes 5 m or more, from the nest before taking flight.

Brooding

Nest attendance decreases during the nestling period from an average of 52.1% (n = 2 nests) in the early-nestling period (days 0–3), to 46.1% (n = 5 nests) in the mid-nestling period (days 4–6), and 10.2% in the late-nestling period (days 7–10) (n = 3 nests). In the early-nestling period, the mean duration of a brooding on-bout is 11.31 min ± 7.85 SD (range 1–37 min; n = 42 on-bouts ), and the mean recess duration is 10.40 min ± 7.93 SD (range 1–43 min; n = 42 recess bouts). In the mid-nestling period, the mean on-bout and recess duration are 10.22 min ± 8.85 SD (range 2–40 min; n = 76 on-bouts) and 12.75 min ± 12.87 SD (range 1–87 min; n = 72 recess bouts), respectively. Although brooding still occurs in the late-nestling period, it mainly involves shading the nestlings. The mean brooding on-bout duration during this period is short (7.5 min ± 7.95 SD, median = 31 min, range 1–31 min; n = 12 on-bouts), but the nestlings are left unattended for longer periods (52.8 min ± 64.48 SD, median = 26 min, range 1–187 min; n = 15 recess bouts) (DE, unpublished data).

Feeding

Males tend to deliver prey at a higher rate, mainly because females are also occupied with brooding duties. Nestlings are fed invertebrates exclusively and are provisioned with the same types of prey consumed by adults (See Diet and Foraging). In the early stages of the nestling period, prey items are usually smaller and soft-bodied, e.g., lepidopteran larvae and adults, termites (Isoptera), and spiders (Arachnida), but in the later stages, grasshoppers (Orthoptera) begin to dominate (DE, unpublished data). Prey with potentially hazardous body parts, e.g., the legs of grasshoppers, are prepared by removing the legs before they are given to the nestlings. Prey such as grasshoppers are either decapitated or, if sufficiently small, presented head first. Prey loading is common, and if more than one prey item is brought to the nest, the prey is usually shared among the brood. If nestlings do not take the prey on offer, the adult will consume it. If a male arrives with food, the female will briefly leave the nest to allow the male to feed the nestlings before returning to the nest to resume brooding (5).

Males deliver prey at a relatively constant rate throughout the nestling period: 3.88 deliveries/h in the early nestling period compared to 4.91 deliveries/h in the late nestling period. The delivery rate of females shows a substantial increase as the nestling period progresses and brooding is reduced: 2.50 deliveries/h in the early nestling period compared to 4.25 deliveries/h in the late nestling period (DE, unpublished data).

Nest Sanitation

Both sexes perform nest sanitation duties. Fecal sacs of nestlings are initially consumed up to about day seven, or are removed and disposed of between 20 and 80 m from the nest. Nestlings will position themselves by defecating at the rim of the nest. There may be an accumulation of droppings in the latter stages of the nestling period , but, in general, the parents remove fecal sacs at every nest visit (47, 6, 5).

Carrying of Eggs or Young

No evidence for either.

Cooperative Breeding

Not known to occur.

Brood Parasitism by Other Species

Not known to occur.

Fledgling Stage

Departure from the Nest

The young fledge at 11–12 d (6, 5).

Association with Parents or Other Young

Fledglings are usually provisioned for 2–3 weeks after fledging (5). Occasionally, the parents can be seen feeding a fledgling up to at least four weeks after leaving the nest when they are also provisioning the nestlings of a subsequent brood (5). The limited data on post-juvenile dispersal suggest the young remain in their natal territory for at least 6–8 weeks after fledging, sometimes as long as 16 weeks. Although both sexes provision the fledglings, most provisioning is by males as females may start constructing a new nest within 2–3 d after the nestlings fledge in the peak breeding season.

Ability to get Around, Feed, and Care for Self

Upon fledging, the young are mobile on the ground and either shuffle or jump short distances but can fly short distances about 10 d after fledging. Fledglings tend to hide amongst vegetation, remaining motionless with their eyes open as slits and relying on their cryptic plumage to escape detection.

Immature Stage

The limited data about post-juvenile dispersal suggest juveniles disperse during their post-juvenile molt. A juvenile that fledged in early November 2003 was resighted 380 m from its nest site four months, 25 d later, while another individual was seen 800 m from its natal territory when it was seven months old (5, 45). Although two additional records suggest limited dispersal by juveniles, e.g., 1,200 m from its natal territory at age 14 months and 3,400 m when the individual was 18 months old (5), none of 23 color-banded nestlings were recorded as adults in the Polokwane Game Reserve; this suggests juveniles disperse further than adult females.

Recommended Citation

Engelbrecht, G. D. (2023). Short-clawed Lark (Certhilauda chuana), version 2.0. In Birds of the World (D. G. Allan, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bow.shclar1.02
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