Comparative Life History of the South Temperate Cape Penduline Tit (Anthoscopus Minutus) and North Temperate Remizidae Species
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J Ornithol DOI 10.1007/s10336-016-1417-4 ORIGINAL ARTICLE Comparative life history of the south temperate Cape Penduline Tit (Anthoscopus minutus) and north temperate Remizidae species 1,2 1 1 Penn Lloyd • Bernhard D. Frauenknecht • Morne´ A. du Plessis • Thomas E. Martin3 Received: 19 June 2016 / Revised: 22 October 2016 / Accepted: 14 November 2016 Ó Dt. Ornithologen-Gesellschaft e.V. 2016 Abstract We studied the breeding biology of the south parental nestling care. Consequently, in comparison to the temperate Cape Penduline Tit (Anthoscopus minutus)in other two species, the Cape Penduline Tit exhibits greater order to compare its life history traits with those of related nest attentiveness during incubation, a similar per-nestling north temperate members of the family Remizidae, namely feeding rate and greater post-fledging survival. Its rela- the Eurasian Penduline Tit (Remiz pendulinus) and the tively large clutch size, high parental investment and Verdin (Auriparus flaviceps). We used this comparison to associated high adult mortality in a less seasonal environ- test key predictions of three hypotheses thought to explain ment are consistent with key predictions of the adult latitudinal variation in life histories among bird species— mortality hypothesis but not with key predictions of the the seasonality and food limitation hypothesis, nest pre- seasonality and food limitation hypothesis in explaining dation hypothesis and adult mortality hypothesis. Contrary life history variation among Remizidae species. These to the general pattern of smaller clutch size and lower adult results add to a growing body of evidence of the impor- mortality among south-temperate birds living in less sea- tance of age-specific mortality in shaping life history sonal environments, the Cape Penduline Tit has a clutch evolution. size larger than that of the Verdin and similar to that of the Eurasian Penduline Tit, and higher adult mortality than Keywords Cooperative breeding Á Nest predation Á both of the other two species. The most notable difference Reproductive success Á Life history traits between the Cape Penduline Tit and the two other species is in parental behavioural strategy, with the former Zusammenfassung exhibiting bi-parental care at all stages of nesting together with facultative cooperative breeding, whereas the Eur- Vergleichende Lebensgeschichte der in der su¨dlichen asian Penduline Tit has uni-parental care and the Verdin gema¨ßigten Zone vorkommenden Kapbeutelmeise has a combination of female-only incubation but bi- (Anthoscopus minutus) und den in der no¨rdlichen gema¨ßigten Zone vorkommenden Beutelmeisen (Remizidae) Communicated by F. Bairlein. & Penn Lloyd Wir untersuchten die Brutbiologie der in der su¨dlichen [email protected] gema¨ßigten Zone vorkommenden Kapbeutelmeise 1 (Anthoscopus minutus), um ihre Lebensgeschichte (life DST/NRF Centre of Excellence, Percy FitzPatrick Institute, history) mit denen verwandter, in der no¨rdlichen University of Cape Town, Private Bag X3, Rondebosch 7701, South Africa gema¨ßigten Zone vorkommenden Mitgliedern der Familie Remizidae, na¨mlich Beutelmeise (Remiz pendulinus) und 2 Present Address: Biodiversity Assessment and Management Pty Ltd, P.O. Box 1376, Cleveland 4163, Australia Goldko¨pfchen (Auriparus flaviceps), vergleichen zu ko¨nnen. Wir haben diesen Vergleich gemacht, um die 3 U. S. Geological Survey Montana Cooperative Wildlife Research Unit, University of Montana, Missoula, MT 59812, Hauptvorhersagen dreier Hypothesen zu testen, welche die USA mit dem Breitengrad zusammenha¨ngende Variation in der 123 J Ornithol Lebensgeschichte von Vogelarten zu erkla¨ren versuchen, incidence of cooperative breeding (Jetz and Rubenstein die Saisonabha¨ngigkeits- und Nahrungslimitations- 2011). Yet, the reasons for these latitudinal differences Hypothese, die Nestpra¨dations-Hypothese und die remain unclear, and further comparisons can benefit Altvogelmortalita¨ts-Hypothese. U¨ blicherweise haben clarification. Arten der su¨dlichen gema¨ßigten Zone, die in weniger Differences in the life history of north temperate and saisonalen Umwelten leben, kleinere Gelege und niedrigere south temperate birds have been explained on the basis of Altvogelmortalita¨t. Entgegengesetzt zu diesem Muster hat three main hypotheses, namely the seasonality and food die Kapbeutelmeise jedoch gro¨ßere Gelege als das limitation hypothesis, adult mortality hypothesis and the Goldko¨pfchen, eine a¨hnliche Gelegegro¨ße wie die nest predation hypothesis. The seasonality and food limi- Beutelmeise und ho¨here Altvogelmortalita¨t als beide tation hypothesis proposes that populations are regulated Arten. Die Kapbeutelmeise unterscheidet sich am during the non-breeding (typically winter) season when sta¨rksten hinsichtlich ihrer Brutpflegestrategie—sie weist food resources are scarcer. In less seasonal environments, in allen Stadien der Brut biparentale Brutpflege nebst such as those of the south temperate zone (Chown et al. fakultativem kooperativem Bru¨ten auf, wohingegen bei der 2004; Lloyd et al. 2014), a lower differential in food Beutelmeise uniparentale Brutpflege und beim availability between non-breeding and breeding seasons Goldko¨pfchen eine Kombination aus alleiniger Bebru¨tung results in greater density-dependent competition and durch das Weibchen und biparentaler Nestlingsfu¨rsorge reduced food availability during the breeding season vorkommen. Folglich verbringt die Kapbeutelmeise (Ashmole 1963; Ricklefs 1980). Reduced food availability wa¨hrend der Bebru¨tung mehr Zeit auf dem Nest und hat is predicted to result in smaller clutch size, reduced incu- eine a¨hnliche Fu¨tterungsrate pro Nestling und besseres bation attentiveness, reduced per-nestling feeding rates, U¨ berleben nach dem Ausfliegen. Die relativ großen greater brood reduction due to starvation, longer develop- Gelege, das hohe Elterninvestment und die damit ment periods and longer re-nesting intervals (Gill and verbundene ho¨here Altvogelmortalita¨t bei der Haggerty 2012). The adult mortality hypothesis proposes Kapbeutelmeise in einer weniger saisonalen Umwelt that the lower annual adult mortality typical of south stimmen mit den Hauptvorhersagen der temperate birds selects for reduced total parental effort in Altvogelmortalita¨ts-Hypothese u¨berein, jedoch nicht mit reproduction but increased parental investment per off- denen der Saisonabha¨ngigkeits- und Nahrungslimitations- spring to enhance offspring survival (Martin Hypothese, wenn es darum geht, Variation in der 1996, 2002, 2014, 2015). The adult mortality hypothesis Lebensgeschichte zwischen verschiedenen Arten der predicts smaller clutch size, reduced incubation attentive- Familie Remizidae zu erkla¨ren. Diese Ergebnisse liefern ness, greater per-nestling feeding rates, lower brood weitere Belege fu¨r die Bedeutung altersspezifischer reduction due to less starvation, longer development peri- Mortalita¨t bei der Evolution der Lebensgeschichte von ods and longer re-nesting intervals among species with Arten. lower annual adult mortality (Gill and Haggerty 2012; Martin 2002, 2014, 2015). The lower annual adult mor- tality of south temperate species may be a consequence of lower seasonality and milder minimum winter tempera- Introduction tures resulting from the moderating influence on climate of a greater ratio of ocean to land in the south temperate zone Comparison of the ecologies of closely related species by comparison with the north temperate zone (Chown et al. occupying different geographical regions and environments 2004; Lloyd et al. 2014). Finally, at the same time, nest can provide valuable insights into the evolution of life predation can interact with adult mortality as part of age- history and behaviour (Martin 1996; Bennett and Owens specific mortality hypotheses (Law 1979; Michod 1979; 2002). Such studies have established that birds of the south Martin 2002, 2014, 2015; Martin et al. 2015). The temperate zone tend to lay smaller clutches (Martin et al. increased nest predation risk typical of south temperate 2000, 2006; Llambı´as et al. 2015) and have longer devel- birds can favor reduced clutch size, lower nestling feeding opment periods (Martin et al. 2007; Llambı´as et al. 2015) rates and faster development (Skutch 1949; Martin et al. and lower adult (Lloyd 2008; Lloyd et al. 2014) and 2000; Martin 2002, 2014, 2015). We examine these three fledgling mortality (Lloyd and Martin 2016) than related hypotheses based on comparisons of three songbird species species in the north temperate zone. Behaviour of south in the family Remizidae. temperate species also tends to differ, with south temperate In this study, we describe the breeding biology of the species showing reduced incubation attentiveness (Chal- little-studied south temperate Cape Penduline Tit (Antho- foun and Martin 2007; Martin et al. 2007), longer post- scopus minutus) to enable a comparison of its life history fledging care (Russell et al. 2004) and exhibiting a higher traits with those of the related and well-studied north 123 J Ornithol temperate Eurasian Penduline Tit (Remiz pendulinus) and (Sony Corp., Tokyo, Japan) mounted on tripods. Nests the Verdin (Auriparus flaviceps), all small insectivores. were video-taped at two stages during incubation, namely One of Africa’s smallest birds, the Cape Penduline Tit early incubation (between 2 and 5 days after the last egg of inhabits semi-arid to arid shrublands and savannas in the clutch was laid) and late incubation (between 9 and southern Africa (Dean 2005). 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