Незамысловатая песня помогает птицам избегать хищников

Рис. 1. Тусклая зарничка (Phylloscopus humei) — пеночка с чрезвычайно простой песней, обитающая в горах Центральной Азии. Фото с сайта oiseaux.net

Тусклая зарничка (Phylloscopus (inornatus) humei) — маленькая, неброско окрашенная птица семейства славковых, живущая в горах Центральной Азии. Ее пение — короткий, негромкий и однообразный свист — столь же непритязательно, как и внешний облик. А ведь песня — один из главных признаков, отвечающих за привлекательность самца. Почему же ее эволюция ограничилась такими скромными результатами? Может ли столь простой сигнал служить показателем качества самца? Китайским орнитологам удалось подтвердить, что самки предпочитают супругов с максимально упрощенной песней — возможно, потому, что такая песня меньше привлекает хищников, разоряющих гнезда. В то же время для оценки физического состояния партнера самки используют альтернативные визуальные признаки.

Пение соловья нетрудно отличить от песни скворца, трели жаворонка — от щебета ласточки. Песня, красивый и сложный элемент птичьего «языка», играет важнейшую роль в коммуникации, но имеет ли значение ее конкретный «рисунок»? На первый взгляд может показаться, что, коль скоро любая песня в конечном итоге обеспечивает свои функции — а это, прежде всего, привлечение противоположного пола и охрана территории, — то любая «мелодия» одинаково хороша. Тем более что она находится под влиянием полового отбора, исход которого зависит от пристрастий самок. Впрочем, ссылка на половой отбор мало что объясняет: совершенно непонятно, почему в сходных, казалось бы, ситуациях, он подчас приводит к противоположным результатам. Как писал Пушкин, «В лесах, во мраке ночи праздной / Весны певец разнообразный / Урчит, и свищет, и гремит; / Но бестолковая кукушка, / Самолюбивая болтушка, / Одно куку свое твердит». Конечно, соловей и кукушка — неродственные птицы, которые ведут совершенно разный образ жизни, поэтому существенные отличия в их «вокальной программе» совсем не удивительны. Но подобные контрасты нередко демонстрируют и близкородственные виды.

Если обыкновенный соловей (Luscinia luscinia) — один из лучших певцов, то соловей-свистун (Luscinia sibilans), как указывает само его название, исполняет лишь простую однообразную трель. Песня пеночек (Phylloscopus) тоже не слишком сложна, если сравнивать их с другими представителями семейства, например камышевками (Acrocephalus). Но тусклая зарничка, или тусклая пеночка (Ph. humei), своим минимализмом превосходит даже незамысловатые песни пеночки-трещотки и таловки. Именно этот вид и стал объектом изучения коллектива китайских орнитологов, которые задались вопросом: как самцам тусклой зарнички удается обходиться столь примитивными сигналами? Их статья озаглавлена Can simple songs express useful signals for mate choice? («Могут ли простые песни служить индикаторам для выбора партнера?»).

Окраска оперения тусклой пеночки неброская, серая с желто-зелеными оттенками; выделяются лишь светлая бровь и полосы на крыльях (рис. 1). Часть систематиков считает ее подвидом пеночки-зарнички (Ph. inornatus), широко распространенной в сибирской тайге. Ареал тусклой пеночки ограничен горами Центральной Азии — Гималаями, Тянь-Шанем, Алтаем и Саянами. Внешне два вида почти неразличимы, но если песня зарнички — это трехсоставная фраза (ее можно передать как «тсьИу-тьиИсси-тсииии»), то тусклая зарничка исполняет лишь короткий двусложный сигнал («вьиси-вьиси» или «тьиу-тьиу»), который перемежается тихим хрипловатым свистом. Изолированный подвид Ph. h. mandellii, обитающий на востоке Гималаев, и вовсе исполняет лишь этот свист (рис. 2), но при этом отличается необычным для пеночек разнообразием позывок.

Незамысловатая песня помогает птицам избегать хищников

Рис. 2. Пение восточного подвида тусклой зарнички (Phylloscopus humei mandellii) в графическом представлении. Вверху — несколько песен, внизу — увеличенный фрагмент с одной песней. Вверху каждого рисунка — осциллограмма (динамика амплитуды звука), внизу — сонограмма (динамика частоты звука в кГц). По оси абсцисс — время (в секундах). Рисунок из обсуждаемой статьи в Avian Research

Авторы отмечают, что подобные примитивные песни нечасто попадают в круг внимания исследователей, хотя их изучение чрезвычайно интересно для раскрытия механизмов полового отбора. Как правило, самки предпочитают самцов с продолжительной и разнообразной песней. Вокальная одаренность служит косвенным показателем здоровья и силы будущего «отца семейства», ведь пение — весьма энергозатратный процесс, а запоминание новых элементов и последовательностей требует незаурядной памяти. Иногда связь между особенностями вокализации и признаками самца совсем неочевидна: к примеру, у лугового чекана быстрый темп песни свидетельствует о большем усердии при охране гнездового участка, а позднее — выкармливании птенцов. Впрочем, о качестве самца может сигнализировать не только песня, но и внешние признаки, такие как окраска или размер. Скажем, у больших синиц на способности самца к защите выводка указывает ширина черной полосы на брюхе. Бывают и более специфические требования: например, самки камышницы предпочитают жирных самцов, которые более эффективно исполняют роль «инкубатора» для кладки.

Однако на пути безграничного развития всевозможных дарований в угоду половому отбору встает «обычный» естественный отбор. В самом деле, если самец будет отдавать все свои силы и способности пению, то это неизбежно приведет к истощению, сокращению продолжительности жизни, а то и вовсе гибели неосторожного певца в когтях хищника. Тем более что на плечи сильного пола нередко ложатся и заботы о птенцах. По-видимому, именно этот «противовес» и устанавливает баланс между простотой и сложностью в песне. Почему же в случае тусклой зарнички равновесие так сильно сместилось в сторону простоты?

Для ответа на этот вопрос авторы изучили социальную жизнь тусклой теньковки (подвид mandellii) в заповеднике Ляньхуашань (провинция Ганьсу, Восточный Китай). Они выявили предпочтения самок при выборе партнера и оценили последствия этого выбора для выживаемости птенцов. Исследуемая популяция пеночек обитала в смешанных лесах, состоящих из ели, сосны, березы и ивы, на высоте примерно 2850 м. Самцов, пойманных паутинными сетями, — всего 55 особей — пометили цветными кольцами для индивидуального распознавания, а также измерили несколько ключевых внешних признаков, которые теоретически могут оцениваться самками. Это длина клюва, крыла, хвоста и цевки, характеризующие размер тела; длина и ширина «украшений» оперения — бровей и полос на крыльях. Пение самцов записывали на диктофон, и затем анализировали длину отдельных песен, интервалы между ними, максимальную амплитуду звука, время от начала песни до ее достижения, а также несколько характеристик частоты звука. Не остались без внимания и самки: орнитологи выслеживали их во время постройки гнезд (конец мая — начало июня), измеряли размер и массу яиц. На 12-й день после вылупления, незадолго до вылета, проводили стандартные измерения размеров птенцов.

Предположив, что наиболее привлекательные особи образуют пару раньше других, авторы использовали дату начала кладки в качестве показателя привлекательности того или иного самца. Начав размножение раньше других, родители в среднем выводили больше птенцов, а сами птенцы были крупнее, чем при поздней кладке (рис. 3). По-видимому, это связано с лучшими кормовыми условиями в начале сезона. А вот ожидаемой связи между размерами яиц и сроками гнездования выявить не удалось: значит, самки вкладывают одинаковое количество ресурсов и в ранние, и в поздние, менее «перспективные» кладки.

Незамысловатая песня помогает птицам избегать хищников

Рис. 3. Корреляции между датой начала кладки (D, дни от самой ранней кладки в конкретный год исследований, слева), объемом яиц (в см3, справа) и длиной клюва птенцов примерно за два дня до вылета из гнезда. Более крупными были птенцы ранних кладок, вылупившиеся из крупных яиц. При этом сам размер яиц в ранних и поздних кладках в среднем был одинаковым. Рисунок из обсуждаемой статьи в Avian Research

Дата начала кладки оказалась связанной как с размерами тела самца, так и с длительностью его песни (рис. 4). Более успешными у самок были крупные самцы, исполняющие короткую песню с быстро нарастающей амплитудой. При этом величина белых полос на крыле коррелировала с размером тела — а значит, по этим маркерам самки могут легко оценить качество их обладателя. Такая возможность ранее уже получила экспериментальное подтверждение в работе К. Марчетти на пеночке-зарничке: искусственное уменьшение ширины полос резко снижало привлекательность самцов, несмотря на достоинства их пения (K. Marchetti, 2002. The evolution of multiple male traits in the yellow-browed leaf warbler).

Незамысловатая песня помогает птицам избегать хищников

Рис. 4. Корреляции между датой начала кладки (D, дни от самой ранней кладки в конкретный год исследований), длиной крыла (в см, слева) и продолжительностью песни до достижения максимальной амплитуды (DSM, в секундах, справа). Наиболее привлекательными для самок были крупные самцы с короткой песней — они образовывали пару и начинали кладку раньше других. Рисунок из обсуждаемой статьи в Avian Research

В выборе крупных «супругов» ничего удивительного нет, ведь размер — надежный показатель физиологического состояния особи. А вот предпочтение короткой песни явно не укладывается в типичную закономерность, когда большей популярностью пользуются длинные и сложные сигналы. Иногда простоту песни может компенсировать яркость окраски, как, например, в семействе лесных певунов — сходных с пеночками и славками североамериканских птиц. Однако это далеко не универсальное правило: скажем, варакушка — самый яркоокрашенный европейский соловей — исполняет не менее сложную песню, нежели невзрачный обыкновенный соловей. В случае с тусклой пеночкой ситуация обратная: она уступает другим представителям рода как по яркости окраски, так и по вокальному разнообразию. Чем же привлекательны для самок упрощенные песни? Авторы исследования связывают это с избеганием хищников — тусклая пеночка строит гнезда на земле, где они легкодоступны для хищников, а длинная сложная песня может привлечь их внимание. Впрочем, это объяснение весьма уязвимо для критики. В конце концов, большинство видов пеночек гнездится на земле, однако ни у кого из них упрощение песни не зашло так далеко. И вряд ли тусклая зарничка испытывает какой-то особый пресс хищников: во всяком случае, никаких свидетельств в пользу этого авторы не приводят.

Другое предположение авторов выглядит более убедительно. Самцы многих птиц, например бурой пеночки (Ph. fuscatus), исполняют два типа песни: длинный и «красивый» предназначен для привлечения самки, а короткий используется в целях охраны территории. При исполнении короткой песни самцу легче услышать пение соседей и соответствующим образом реагировать на них. Поэтому не исключено, что самки тусклой пеночки отдают предпочтение именно таким успешным защитникам своего участка.

Упрощенную песню можно связать и с участием самцов в заботе о потомстве: ведь хорошее физиологическое состояние самца еще не гарантирует, что он будет вкладывать свои силы в выкармливание птенцов. Так, детальные исследования В. Форстмайера с соавторами показали, что некрупные и слабые самки бурой пеночки образуют пары с такими же «второразрядными» самцами, которые компенсируют недостаток физической мощи усердием в «домашних хлопотах» (W. Forstmeier et al., 2002. A novel song parameter correlates with extra-pair paternity and reflects male longevity).

Правда, это не мешает самкам встречаться и с другими «кавалерами», так что слабому самцу обычно приходится заботиться и о чужих отпрысках. Тем не менее его стратегия вполне успешно сосуществует со стратегией доминантов, которые зачастую вообще не помогают в выкармливании. Вполне возможно, что в суровых условиях гималайских горных лесов без участия самца в кормлении птенцов не обойтись — и поэтому самки тусклой зарнички стали отдавать предпочтение партнерам, которые не растрачивают слишком много сил на брачные демонстрации. Впрочем, Лью и его коллеги не оценивали процент внебрачного отцовства, так что их результаты говорят об успехе самцов лишь как социальных партнеров. Не исключено, что при внебрачных спариваниях более привлекательными оказываются самцы с какими-то другими признаками. Излишне добавлять, что в брачном поведении воробьиных птиц едва ли присутствуют какие-либо персонализированные отношения и привязанности.

Стоит отметить и то, длинная и сложная песня — не самоцель, а лишь показатель энергетических расходов самца. Может быть, в случае тусклой пеночки короткий стремительный свист требует от самцов больших затрат, чем более долгий? К сожалению, этот вопрос пока что остался без ответа. Однако авторы установили, что интервал между песнями не влиял на мужскую привлекательность — значит, самки «ценят» короткую песню саму по себе, даже если она исполняется с большими интервалами. А вот у пеночки-зарнички, как показала упомянутая выше работа Марчетти, такая связь имеется: это свидетельствует о разнообразии путей полового отбора даже в пределах близкородственной группы видов.

Итак, на вопрос, сформулированный в названии статьи, авторы дают положительный ответ. Простые сигналы вполне справляются со своей задачей и успешно помогают самкам выбрать пару, хотя почему они предпочитают именно такой сигнал — еще не до конца понятно. Кроме того, самки дополняют свою оценку визуальными характеристиками, и для этого им, опять-таки, вполне достаточно весьма незначительных — с человеческой точки зрения — различий. Возвращаясь к вопросу о том, имеет ли функциональное значение структура песни, уместно провести аналогию с музыкой. Конкретная последовательность нот в музыкальном произведении может бесконечно варьировать, и, в то же время, мы легко отличим полонез от марша, печальную музыку — от веселой. Подобно этому, общие характеристики птичьей песни, такие как громкость, темп или разнообразие, тоже подчиняются определенным правилам, которые диктует их социальная функция. При этом в деталях песни могут быть сколь угодно разнообразными — как в сторону усложнения, так и в сторону радикального упрощения.

Источник: elementy.ru

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