Seasonal Trends in Summer Diet of the Lapland Longspur Near Barrow, Alaska
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Condov, SO:295301 0 The Cooper Ornithological Society 1978 SEASONAL TRENDS IN SUMMER DIET OF THE LAPLAND LONGSPUR NEAR BARROW, ALASKA THOMAS W. CUSTER AND FRANK A. PITELKA The Lapland Longspur (Calcarius lapponicus), spur that could be used in an energetic model near Barrow, Alaska, is the only major pas- to calculate number of prey items required serine in a terrestrial insectivorous guild which per day to sustain existence (Custer 1974). mainly consists of shorebirds. The guild in- Here we examine seasonal trends in the diet cludes four sandpipers of the genus Calidris and then compare them with existing data (alpinu, melanotos, pusilla, and bairdii), the from other members of the avian tundra com- American Golden Plover ( Pluvialis dominica ), munity. Information on breeding density, re- the Ruddy Turnstone ( Arenaria interpres), productive success, and adult survival for and the Red Phalarope (Phalaropus full- longspurs at Barrow is presented elsewhere carius) . Other species that rely on arthropods ( Custer and Pitelka 1977). but are irregularly or only locally present in- clude the Snow Bunting (Plectrophenax niva- METHODS lis) and eight additional shorebird species Information on longspur diet was obtained in two (Pitelka 1974). Jaegers, gulls, and waterfowl ways between 28 May and 20 August from years consume arthropods but their total impact is 1969, 1971, 1972, and 1973. First, male, female, and much less than the other species (MacLean juvenile longspurs were collected within 10 km of 1969 ) . the Naval Arctic Research Laboratory, Barrow, Alaska, and as soon as possible (usually within 5 min) The diversity of potential habitats and prey a solution of 10% formaldehyde was injected into the for insectivores is extremely limited at Barrow stomach using a syringe inserted into the throat. (Holmes 1966). The terrain is a mosaic of The stomachs and esophagi of these birds were then poorly drained lowland and variably drained removed and examined in the laboratory. Second, upland tundra broken by small ponds, lakes specimens of longspurs were obtained from lemming snap-trap lines (see Ritelka 1973). Due to post- and streams. The predominant vegetation is mortem decomposition of stomach contents in the grasses and prostrate sedges underlain with snap-trap specimens, only food items in the esophagi mosses and lichens (Britton 1957, Wiggins were removed and examined. and Thomas 1962). Two families of Diptera The categories of prey identified in the food sam- (Chironomidae and Tipulidae) account for ples are listed in Table 1. The number of each prey type was determined either from whole individuals more than 80% of the insect biomass (Holmes or the number of specific parts present in each 1966). The most conspicuous feature of the stomach. The lengths of larvae were measured when insect fauna near Barrow is the abundance of possible. adult dipterans during July. Members of this Freshly collected living specimens of arthropods were frozen immediately and shipped frozen to order overwinter as larvae and, at least in the Berkeley, California, for analysis. The specimens larger species, this stage lasts several years were dried in an oven at 70°C for 48 h and then (MacLean 1973). Those that pupate in a weighed on a Mettler balance to the nearest 0.01 mg given year emerge within a two-week period (Table 1). The mean weight of a crane fly (Tip& in early July ( MacLean and Pitelka 1971) . A carinifrons) and of chironomid larvae, large numbers of which are eaten by longspurs, was estimated by detailed list of the arthropods at Barrow is using available length-weight regressions and aver- provided by Hurd ( 1958). aging the weights for the lengths actually taken by Two studies concerning the shorebird com- longspurs. munity at Barrow demonstrate that four major Samples of adult and juvenile longspurs were Calidris species are strikingly similar in diet pooled in the analysis of diet. This was necessary because of small sample size and missing age in- (Holmes and Pitelka 1968) and overlap wide- formation for several samples. Combining age groups ly in habitat usage (MacLean 1969). Com- does not appear to bias the data since juveniles were petition among coexisting sandpipers is sug- not collected prior to 1 July, and comparisons be- gested by differences in bill lengths and other tween adult and juvenile longspurs for the periods l-10 July and 11-20 July indicate that the diets are features of bill structure and by the observa- similar. Most of the samples from l-10 August were tion that there are only four consistently abun- of unknown age composition and all samples from dant Calidris species at Barrow. 11-20 August consisted only of juveniles. The dietary information is presented in two ways: The main objective of this study was to pro- ( 1) percent of diet by number of all food items pres- vide dietary information on the Lapland Long- ent in all stomachs and esophagi (enumeration meth- 12951 296 THOMAS W. CUSTER AND FRANK A. PITELKA TABLE 1. Mean weight and correction factor for prey items taken by the Lapland Longspur. Mean dry Co,;~&O Prey item weight ( mg) Collembola 0.06 5.0 Saldidae (immature) 1.57” 5.0b b Saldidae (adult) 3.05 Eh mLARVAL AND IMMATURE ARTHROPODS _ z.: 0 Carabidae ( larvae ) 2.60 DSEEDS Carabidae ( adult ) 5.90 3:9 A! o_ , I I I , , , , _ Staphylinidae ( adult ) 1.20” z$ Chrysomelidae ( larvae ) 9.38 Chrysomelidae (adult) 8.70 3:9b Tip& (larvae) 10.66 ’ 5.0 Male Tipula ( adult) 7.39 4.3 Female Tip& ( adult ) 21.63 4.3 Pedicia ( adult ) 1.00” 6.5 Chironomidae ( larvae ) o.17d 7.5 Other large Nematocera ( adult ) 0.93 5.0” 88 00 Small Nematocera ( adult ) 0.20 5.0” Muscidae (larvae) 1.82 5.0 Muscidae (pupae) 1.90 5.0b Large Brachycera (adult) 3.28 5.0 Small Brachycera ( adult ) 0.23 5.0 Tenthredinidae ( larvae ) 6.05 5.0” Hymenoptera ( adult ) 0.20 5.0b Arachnida 1.41 ;$ Nematoda 0.20” ~uzula confusum ( seed ) 0.28 l:ob Cerustium sp. (seed) 0.26 l.Ob JUNE ’ JULY ’ AUGUST Calamagrostris or Luzula (seed) 0.16 l.Ob FIGURE 1. Diet of the Lapland Longspur in four Unidentified seed 0.20” l.Ob summers (1969, 1971, 1972, and 1973) at Barrow, Moss capsule 0.32 l.Ob expressed in: (A) percent by number; (B) corrected n Data from Holmes ( 1966). percent by number; and ( C ) corrected percent by bEstimated using similar species. dry weight. Numbers above bars refer to the total (1Data provided by L. Clement (per% comm.). d Data provided by D. Bierle (per% comm. ). stomachs and esophagi examined from successive e Estimate used in calculation. time intervals. spurs and shorebirds included five immature cate- od of Hartley 1948); and (2) percent of diet by dry gories (Tip& larvae, chironomid larvae, muscid lar- weight, calculated by multiplying the number of vae, muscid pupae, and coleopterous larvae) and items of each prey category by its respective dry seven adult categories (Tip&, other Diptera, Hy- weight (Table 1) and then expressing these values menoptera, Carabidae, Staphylinidae, Arachnida, and as a percent of total dry weight. Collembola). The stomach samples were corrected for differ- Percent overlap between longspurs and shorebirds ential digestion of prey items following the proce- was calculated by following the method of Holmes dure of Custer and Pitelka ( 1975). In this prpce- and Pitelka (1968), in which the minimal percent dure, seeds are held constant and other prey items of each food resource shared by two species is are multiplied by their relative disappearance time summed. Thus, two species with identical resource in relation to seeds. The correction factor used for usage would have an overlap of 100 and two with each food item is listed in Table 1. Esophageal sam- completely different resource usage would have an ples were not corrected for differential digestion. overlap of zero. Diets of longspurs can be compared with those of shorebirds only on the basis of numbers of items present. Because shorebirds do not eat seeds and be- RESULTS cause Holmes and Pitelka ( 1968) did not correct for A total of 8,849 prey items were identified differential digestion, we adjusted the longspur data to make them comparable with the shorebird in- from 174 stomachs and 61 esophagi of long- formation. A single correction factor of 5.0 was used, spurs. Figure 1A illustrates the percent com- as an average of the ratio of time over which seeds position by number of prey items divided into versus arthropods persist in the stomach (Custer three basic categories (seeds, immature ar- and Pitelka 1975). Thus, values for seeds from long- spur stomachs were divided by five and percent rep- thropods, and adult arthropods) in successive resentations were newly obtained. Then, overlaps lo-day intervals of the summer. with the shorebird diets were calculated. In this way, When percent composition of the diet is the arthropod fraction of the longspur diet was ad- corrected for differential digestion rates of justed to bring the numerical intakes of the two kinds various prey types, the significance of seeds of birds onto a comparable basis. The number of prey categories used to calculate overlap between long- drops (Fig. 1B). Percent composition by dry LAPLAND LONGSPUR DIET 297 KEY m ARACHNIDA n Tipula B COLLEMBOLA CHIRONOMIDAE ~Z%IBRACHYCERA gx&y OTHER NEMATOCERA m TENTHREDINIDAE -4 m COLEOPTERA ANC HEMIPTERAT IIII]OTHEFi HYMENOPTERA A LARVAL C LARVAL AND AND IMMATURE ARTHROPODS IMMATURE ARTHROPODS .I :; _[ y. .:. ,;i .I. r.1. A ( 3 I B IID ADULT ARTHROPODS II ADULT ARTHROPODS II I ” A! AUGUST FIGURE 2. Corrected percent composition by number and by dry weight of the arthropod portion of the Lapland Longspur diet divided into major categories of adult and immature arthropods.