У черно-белых ястребов больше птенцов выращивают родители с разной окраской

Рис. 1. Светлая (слева) и темная морфы черно-белого ястреба. Как и для других хищных птиц, для этого вида характерен обратный половой диморфизм: самка (справа) крупнее самца (в данном случае — почти в 1,5 раза). Как самцы, так и самки черно-белого ястреба могут быть и темными, и светлыми, причем это соотношение в пределах конкретной популяции одинаково. Фото с сайта focus-natur.de

В популяциях африканского черно-белого ястреба присутствуют, независимо от пола, птицы темной и светлой морф. Они различаются не только цветом, но также и поведением и/или физиологией. Наличие двух дискретных морфотипов дает уникальную возможность выяснить, как комбинация морф родителей влияет на успех их размножения. Что лучше: разные, взаимно дополняющие родители или одинаковые, не мешающие друг другу своими несхожими привычками? Оказалось, что наибольший репродуктивный успех достигается тогда, когда пара смешанная, то есть когда партнеры относятся к разным морфам. Более детальный анализ позволил выявить и некоторые дополнительные подробности.

Полиморфизм — это, в частности, наличие в популяции двух или более морф. Среди птиц (а это примерно 10 000 видов) полиморфизм известен только у 3–5%. В наибольшей степени он характерен для хищных птиц (отряды ястребообразные Accipitriformes, соколообразные Falconiformes и совообразные Strigiformes). Так, в первых двух отрядах это свойство суммарно характерно для 30% видов (M. K. Fowlie, O. Krüger, 2003. The evolution of plumage polymorphism in birds of prey and owls: The apostatic selection hypothesis revisited). Известно, что разные морфы отличаются не только окраской, но и другими особенностями — например, физиологией и/или поведением. Таким образом, полиморфизм как бы маркирует комплексные различия между индивидуумами. Так что, помимо прочего, это дает ученым уникальную возможность изучать влияние «комбинации» родителей на успех их размножения.

Вид, подвид или морфа?

Часто исследователь сталкивается с ситуацией, когда в одной местности живут очень похожие животные (например, птицы), но стабильно различающиеся окраской. Что это — разные виды или разные морфы одного и того же вида? Решить это исходя только из внешнего вида обычно нельзя. Как правило, для ответа необходимо знать (1) частоты этих «разноокрашенных» птиц в разных частях ареала и (2) скрещиваются ли они свободно или нет. Если ответ на вопрос (2) отрицательный, то это разные, репродуктивно изолированные виды. Если положительный, то необходимо смотреть более детально. Возможны две основные ситуации. В первом случае разноокрашенные птицы занимают один и тот же ареал, хотя относительные частоты в разных местах этого ареала могут различаться. Тогда это разные морфы одного вида. Во втором случае в большинстве мест можно встретить лишь один вариант окраски. И только в местности, где исследователю повезло оказаться, они живут совместно. И это не что иное, как зона гибридизации двух форм. Значит, это не морфы. Но что это за формы — виды или подвиды? Ответить на этот вопрос четко, к сожалению, нельзя. Обычно, если различия между ними комплексные и затрагивают не только окраску, но и поведение, голос и другие особенности, их считают разными видами. А если различия не очень значительные — это подвиды.

А ведь это поле исследований до сих пор остается почти не изученным. Дело тут в практических трудностях. Если качество самцов ученые еще как-то оценивают (например, по его пению или яркости оперения), то с самками, часто скромно окрашенными и не поющими, намного сложнее. Так что наличие двух морф как самцов, так и самок в популяции одного вида дает возможность ответить на один на первый взгляд простой вопрос: что лучше — разные, взаимно дополняющие родители или одинаковые, не мешающие друг другу своими несхожими привычками?

На этот вопрос и попыталась ответить группа южно-африканских орнитологов. Объект их исследования — черно-белый ястреб Accipiter melanoleucus. Он обитает в Африке к югу от Сахары. Существуют две морфы: темная и светлая (рис. 1). Генетически наследование окраски можно аппроксимировать лишь одним локусом, доминантный аллель которого соответствует светлой морфе (A. Amar et al., 2012. Genetic plumage polymorphism in a newly colonized Black Sparrowhawk population: classification, temporal stability and inheritance patterns). Морфы различаются также особенностями приспособления к среде обитания. В частности, в пределах ареала вида частота морф различна и связана с уровнем осадков в данной местности в течение гнездового периода ястребов (рис. 2): темных ястребов больше там, где влажнее. Это соответствует известному правилу Глогера (выдвинутому в 1833 году), которое гласит, что темноокрашенные виды/морфы живут в более влажных и теплых районах, а светлые — в более сухих и холодных.

У черно-белых ястребов больше птенцов выращивают родители с разной окраской

Рис. 2. А — доли темной и светлой морф в разных популяциях на территории ЮАР, цифры указывают на объем выборки. Б — связь доли темной морфы с уровнем осадков в гнездовой период, размер кружков пропорционален размеру выборки. Рисунок из статьи: A. Amar et al., 2014. Clinal variation in the morph ratio of Black Sparrowhawks Accipiter melanoleucus in South Africa and its correlation with environmental variables

В фокусе исследования была популяция черно-белого ястреба, гнездящаяся в Кейптауне (ЮАР). Ее мониторинг осуществляют с 2000 года, но подробные данные, положенные в основу обсуждаемой статьи, были собраны в 2006–2013 гг. На протяжении этих лет ученые отыскивали гнезда ястреба, определяли морфы взрослых птиц и кольцевали птенцов. А затем оценивали выживаемость птенцов (в течение первого года) и отмечали, кто из них затем успешно гнездился (обычно в возрасте около трех лет). В данном случае вероятность загнездиться и дожить до трех лет — это почти одно и то же. Дело в том, что ястребы в Кейптауне оседлые, и родившиеся здесь птицы, если они доживают до репродуктивного возраста, селятся обычно тут же. То есть дисперсия в другие районы выражена в минимальной степени. Так, среднее расстояние между местом рождения и местом первого гнездования было всего 6,65±0,60 км.

В программе MARK была оценена выживаемость птенцов (вероятность дожить до следующего года) в зависимости от морф родителей. В этом анализе участвовало 234 птенца. Оказалось, что этот показатель был выше, если партнеры в паре имели разную окраску, а не одинаковую (рис. 3, А). То есть взаимно дополняющие друг друга родители, разные по окраске, — это наилучший вариант. При этом было не важно, каковы конкретные комбинации. В смешанной паре темным мог быть самец, а могла и самка. А если родители имели одинаковую окраску, она могла быть темной или светлой (рис. 3, Б). Пока точно не ясно, с чем это может быть связано. Однако авторы предполагают, что темные и светлые ястребы различаются охотничьими предпочтениями — что и определяет доминирование темной морфы в более влажном климате. Поэтому если партнеры окрашены различно, они более полно используют имеющиеся на их участке ресурсы и как бы страхуют друг друга. Конечно, это применимо лишь к Кейптауну, где нередки обе морфы, хотя темные заметно преобладают (составляя около 75%).

У черно-белых ястребов больше птенцов выращивают родители с разной окраской

Рис. 3. Выживаемость птенцов в зависимости от морф родителей: А — птицы в паре относятся к разным (смешанная пара) или одинаковым морфам; Б — четыре возможных комбинации, например светлый (С) самец + темная (Т) самка, и т. д. Рисунок из обсуждаемой статьи в Journal of Animal Ecology

На следующем этапе ученые изучали, с какой вероятностью птенец данной пары пополнит гнездовую популяцию. Использовали данные по 187 птенцам, родившихся в период с 2006-го по 2011 гг. Удалось выявить еще одну закономерность. Выяснилось, что вероятность птенца загнездиться зависит от морфы самца, но не самки. Причем это влияние опосредовано временем появления на свет. Наиболее успешными были, с одной стороны, отпрыски темных отцов, родившиеся в начале сезона гнездования. А с другой — потомки светлых самцов, появившиеся в конце этого периода (рис. 4). Это на первый взгляд парадоксальное явление тоже можно объяснить. Дело в том, что у ястребов, как и у других хищных птиц, обязанности между родителями разделены достаточно четко. Более крупная самка насиживает кладку и обогревает птенцов, пока они еще маленькие. А попутно и охраняет гнездо. В обязанности же самца входит снабжать пропитанием сначала самку, а потом и маленьких птенцов. Лишь когда птенцы подрастут, добывать пищу для них начинает и самка. Так что на протяжении заметной части гнездового периода самец играет ключевую роль, обеспечивая семью кормом. В начале периода гнездования (март–июль) погода более влажная. Не исключено, что в это время именно темные самцы ловят добычу успешнее. А в конце (сентябрь–октябрь) становится более сухо, преимущество могут получать уже светлые особи. Август является периодом «переходным».

У черно-белых ястребов больше птенцов выращивают родители с разной окраской

Рис. 4. Ожидаемая вероятность молодых особей загнездиться в зависимости от морфы их отца и месяца вылупления. Рисунок из обсуждаемой статьи в Journal of Animal Ecology

Таким образом, в данном исследовании впервые выявлено комплексное влияние морф родителей на успех их размножения. Очевидно, это, с одной стороны, определяет сосуществование разных морф в одной популяции, а с другой — влияет на различия в частоте их встречаемости в разных частях ареала.

Источник: elementy.ru

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