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Comparative breeding ecology and hybridization of Eastern
and Western Marsh Harriers Circus spilonotus and
C, aeruginosus in the Baikal region of Eastern Siberia
比较繁殖生态学和东欧杂交
西方沼泽鹞鹞和
C，在东西伯利亚的贝加尔湖地区aeruginosus
IGOR V. FEFELOV伊戈费费洛夫
Research Institute of Biology at lrkutsk State University, PO Box 24, 664003 /rkuts,k, Russia
New information on the breeding ecology and inter-breeding of Western Circus
aeruginosus and Eastern C. spilonotus Marsh Harriers in the Baikal region of Eastern Siberia
is presented. Hybridization is relatively common (one of the parents was not ‘pure’ C.
spilonotus in six out of 28 nesting cases) in the Irkutsk area, where nests of a mixed pair
were found with chicks in 1999 and 2000. Eastern Marsh Harriers are more similar to
Western in size, biology and coloration of young birds, but to the Australasian C. approximans
in their displays and mating system, and intermediate between C. aeruginosus and
C. approximans in the plumage coloration of females. The systematic status (species’ or
‘subspecies’) of the Western and Eastern Marsh Harriers depends on the systematic
approach taken, and existing data seem to be inadequate to resolve this. The Siberian
Mongolian zone of contact of these marsh harriers is unique as other marsh harriers of the
world are allopatric. More research on ecology, behaviour and molecular systematics is
necessary to resolve the situation.
生物研究所在lrkutsk州立大学，邮政信箱24，664003 / rkuts，钾，俄罗斯
新的信息的繁殖生态及相互西方马戏育种
aeruginosus和东欧长spilonotus在东西伯利亚的贝加尔湖地区的沼泽鹞
提出。杂交比较普遍（父母一方是不是'纯粹'长
筑巢在28例）spilonotus在伊尔库次克地区，在六项筑巢混合配对
被发现在1999年和2000年小鸡。东方沼泽鹞更类似于
西方的大小，生物学和幼鸟色彩，但对澳长approximans
在他们的显示器和交配系统，以及介于长aeruginosus和
在女性的羽毛色泽长approximans。该系统的状态（物种或
'种'西部和东部沼泽鹞）取决于系统
采取的方式，和现有的数据似乎不足以解决这个问题。西伯利亚
这些沼泽鹞接触蒙区是独一无二的其他沼泽鹞
世界的异域。更多的研究对生态，行为和分子系统学的
要解决这种情况。
The superspecies complex of marsh harriers distributed
through Eurasia, Africa, Australia and some
islands of the Indian and Pacific Oceans comprises five
well-distinguished species including eight distinct
forms (Amadon & Bull 1988, del Hoyo et al. 1994).
However, they are allopatric apart from two races:
Western (Eurasian) Circus (aeruginosus) aeruginosus L.
1758, and Eastern (Striped) C. (a,) spilonotus Kaup
1847, Marsh Harriers. The area of their intergradation
in northwestern Mongolia and southern Central
Siberia (Stepanyan 1990) is, therefore, unique in the
marsh harrier complex and is thus of great interest.
While the ecology of the first species is well-studied in
Europe, little is known about the second. This paper
compares the breeding ecology of these two forms,
how they interrelate in their contact zone in Southern
Siberia, and the systematic implications.
在沼泽鹞superspecies复杂的分布式
通过欧亚，非洲，澳大利亚和一些
印度和太平洋岛屿组成5
包括8个不同的良好杰出的物种
表格（Amadon＆红牛1988年，格雷贝尔德尔奥约等人。1994年）。
然而，他们异域除了两站比赛：
西方（欧亚）马戏（aeruginosus）aeruginosus湖
1758年，和东区（条纹）长（1）spilonotus海龙属
1847年，马师鹞。他们intergradation区
蒙古西北部和南部环
西伯利亚（斯坦波尼扬1990年），因此，独特的
沼泽鹞复杂，因此非常感兴趣。
虽然这是第一个物种的生态良好的研究
欧洲，知之甚少第二。本文
比较了这两种形式的繁殖生态，
如何在他们的相互联系南接触带
西伯利亚和系统的影响。
STUDY AREA AND METHODS
Breeding biology, plumage coloration, food and interbreeding
of marsh harriers were studied during
1989-2000 in the southern Irkutsk Region, in the
Selenga River delta on Lake Baikal (Buryatia
Email: u000438@ic.isu.ru
Republic), at other sites in the Irkutsk Region, and in
1993 in the area of Torey Lakes (Chita Region).
The Selenga delta, the main habitat of C. spilonotus
in Eastern Siberia, has an area of 540 k m 2 . Its upper
part is covered by thickets dominated by willows Salix
spp. The lower part has a large number of lakes and
channels with extensive marshes of Reed Phragmites
australis and other grasses, Broad-leaved Reedmace
Typha latifolia, Sea Sedge Acorns calamus, sedges
Carex spp. etc. Much of it is flooded during high
waters. The water level is minimal in April and May
and maximal in August and September. About 250
k m 2 of the lower and middle parts of the delta are suitable
for marsh harriers. From 150 to 300 pairs of C.
spilonotus nest here; the upper level of 450 pairs previously
published (Fefelov 1996) is probably too high.
A similar landscape occurs in some areas near Irkutsk
and Angarsk, in the valleys of the Irkut and Angara
rivers. However, Reed is largely replaced by Broadleaved
and Narrow-leaved Reedmace ?: angustifolia
here. These wetlands occupy small areas, each of them
being only 1-3 km*.
Forty-four nests of C. spiloiiotus (e.g. 37 in the
Selenga delta), and two nests of a mixed pair C. spilonotus
x C. aeruginosus, were found and observed, and
137 territories of pairs were mapped using binoculars.
@ 2001 British Ornithologists’ Union
588 I.V. Fefelov
Plumage coloration of 87 breeding birds and 144
nestlings was described. About 220 prey samples (pellets,
prey remains, food of nestlings and visual
observations) were collected. Furthermore 70 specimens
of marsh harriers from Siberia and the Far East
were examined in the collections of the Zoological
Museums of Moscow, Irkutsk and Far East
(Vladivostok) State Universities, and of Irkutsk State
Agricultural Academy.
研究范围与方法
繁殖生物学，羽毛色彩，食品和杂交
沼泽鹞的研究中
1989-2000年在南部的伊尔库茨克州，在
色楞格河三角洲的贝加尔湖（布里亚特
电子邮件：u000438@ic.isu.ru
共和国），在伊尔库茨克地区的其他网站，并在
1993年在托列伊湖（赤塔地区）地区。
在色楞格河三角洲，长spilonotus主要栖息地
在东西伯利亚，有540介面积。它的上部
部分覆盖为主的杨柳柳灌丛
属。下部有大量湖泊
渠道广泛的芦苇沼泽芦苇
芦苇和其他草，阔叶Reedmace
宽叶香蒲，海莎草科橡子菖蒲，莎草
苔草。等而在很大程度上是在高洪水
水域。水位是最小的在4月和5月
和最大8月和9月。关于250
三角洲公里的中下层2部分适合
为沼泽鹞。从150至300 C的对
spilonotus巢这儿的四百五十对上层以前
出版（费费洛夫1996年）可能太高。
类似的景观发生在伊尔库茨克附近的一些地区
和安加尔斯克，在伊尔库特和安加拉山谷
河流。但是，里德在很大程度上取代阔叶
窄叶Reedmace？：沙枣
这里。这些湿地占用小区，每个
目前只有1-3公里*.
44个C的spiloiiotus巢（例如，在37
色楞格河三角洲），和两个混合配对筑巢长spilonotus
x长aeruginosus，发现和观察，
137对领土被映射使用双筒望远镜。
@ 2001'英国鸟类学家联盟
588静脉注射费费洛夫
羽毛色泽87繁殖鸟和144
雏鸟被描述。约220猎物样本（颗粒，
猎物遗骸和视觉的雏鸟的食物
观察）收集。另外70个样本
沼泽鹞的从西伯利亚和远东地区
被审查的动物学集合
莫斯科，伊尔库茨克和远东地区博物馆
（海参崴）州立大学和伊尔库茨克州
农业科学院。
分布在沼泽鹞
西伯利亚中部
在长aeruginosus养殖范围向东延伸
而这正spilonotus湖Raikal，延伸
西边的叶尼塞河和西北蒙古
（斯坦波尼扬1990）。这不同于描述
给予格雷贝尔德尔奥约等人。 （1 994），如果这些表格
显示为异域。沼泽鹞分布不均
在叶尼塞河上游河谷（Kokhanovski
1991年），之间的叶尼塞和贝加尔湖地区，
他们更稀少，因为人手短缺
这里最具生产力的湿地，对这些
鸟类而定。
伊尔库次克地区的人口由两鸟
形式。在它的中心，长aeruginosux更频繁
（Mel'nikov 1999年，这项研究），尽管这些数据
不够详细，在南部和东部地区（如靠近
伊尔库茨克州）长spilonotus占主导地位，一些本地网站
在安加拉河流域长aeruginosus（Skalon
1934年）被洪水淹没后布拉茨克创作
水库在20世纪60年代。在对贝加尔湖的西岸，
spilonotus为主体（费费洛夫1996年）;在色楞格河
三角洲所有132筑巢的鸟类观察对被长
spilonotus，以及只有两次（1987和1990年）
是一个成年男性长aeruginosus这里看到的
（图1）。目前在没有C的aemginosus记录
东部贝加尔或进一步向东。
东欧和杂交
西部沼泽鹞
之间的叶尼塞和贝加尔地区，西北
蒙古是intergradation之间的区
长aeruginosus和C. spilonotus，这些都是人口聚居
由表型'纯粹'的两种形式的鸟类以及
图1。在马戏aeruginosus和C spilonotus分布在东西伯利亚南部。 \ \ \，只有射程的aeruginosus / / /，范围
唯一的spilonotus：交叉影线，由aeruginosus和spilonotus居住区;？，足够的数据，十，点上繁殖的aeruginosus
证实了以前或目前的N.点上繁殖的spilonofus证实，糯，点确定
美洲国家aeruginosus和spilonotus育种; -. - 俄罗斯国家边界; -. 。 。 - 俄罗斯的边界行政区域。
0 2001'英国鸟类联盟，宜必思，143，587-592
选育的东部和西部沼泽鹞589
鸟类的羽毛，具有中级（Dement'ev
195 1，斯坦波尼扬1990）。然而，没有什么信息
关于这一点。这两个比赛表现出相当的
个人和时代的变化，这造成重大困难
外地可能的杂交鉴定。
从这些染色的经典描述
表格（Dement'ev 195 1，斯坦波尼扬1990年，韦克
1980年），从我个人的观察，最显着
成人域字符角aeruginosus是
优势的褐色的腹部和排气男性，
缺乏对初选，二级工具和带
尾巴的女性。对于长spilonotus，白色为主
在腹部，通风男性，而女性的
机翼和机尾带羽毛的臀部是
白色或浅。另公布的字符，即
男性blaclush或棕upperparts，是可变的
和不明确。如果我们以此作为基础，对这些字符
识别，没有标本发现，展览的功能
两个长aeruginosus和长spilonotus
上述命名集合。只有一个可能
在莫斯科收集混合成年男性
大学，由雁Przevalsh上采取Lobnor
在中亚湖（无日期），这是大致相同
到C aeruginosus，但有白色喷口和更低
腹的spilonotus（费费洛夫1999年）。此外，成人
与男性之间的长羽毛中间aeruginosus
和C spilonotus收集西北
蒙古（哈拉珠湖）和中国西部
（Yarkend，马扎里沙标签）（施特格曼1937）。一般来说，
目前只有极少数的地区采集标本
其中intergradation很可能发生。
甲'发现'类型的男性长spilonotus，帐户
为在色楞格河三角洲筑巢男性5-10％
（费费洛夫1996年）需要进一步研究，因为它不
羽毛清楚这是否代表着个体差异，
或者是特定于2 - 和3岁的鸟类。他们
不能混合动力车，但是，由于它们都记录
在不同的地区，包括东部的贝加尔，
远东（在上述集合）和
日本（马西等人。1990年）。
良好的自然和栖息场所短缺
人类毁灭的网站在20世纪
在同域区已减少的可能性
相互之间的长aeruginosus育种spilono和C -
一个明显的混合来源的成年男性嵌套
在1993-95在伊尔库茨克附近失事，这可能是相同的
鸟类每年[费费洛夫19991。论颜色
土族。
spilonotus在这些年来。在同一时间从一个
两个典型的长spilonctus对嵌套这里。
嵌套男性长aeruginosus录得近
1995年，1998年，1999年干旱安加尔斯克2000年，成对
女长spilonotus。目前还不清楚是否有
同鸟类每年因为他们没有发现个别
识别功能里里的羽毛。他们的
巢被发现于1999年和2000年。四小鸡出
5，成功孵化蛋，1999年正式（费费洛夫
1999）。五只小鸡（两男三女）
正式于2000年。混合动力幼鸟羽毛
（图2）是非常相似的长spilonotus
（图3）。雏鸟的混合较少暗条纹的
比的（3。spilonotus多数冠。然而，
它们不能被歧视的领域。在1996年和
1997年，长一双spilonotus嵌套在这一领域。
沼泽鹞的30宗已筑巢
安加尔斯克和伊尔库茨克附近失事，录得角二十年
（里亚布采夫和费费洛夫1997年，这一研究）。在其中23
父母双方都长spilonotus四，男性
是长aeruginosus和threc的男性，混合
特征。
繁殖生态
显示
在春，长spilonotus求爱飞行要么
非常相似，非洲沼泽鹞长的
ranivorus多丹1800，和太平洋（澳）
沼泽鹞长approximans欧莱雅eale 1848年，（西蒙斯
1991年），或者是与他们的显示和中间
是C的aeruginosus。在男性和女性长spilonotus，
横向'天舞'及降
upperparts，可以为C，amginosus或C，spilonotus'
它的乳房颜色在C'spilonotusl和
低的排气和在角上午＆ -
nosus。它femalels？）及幼鸟被彩色的长
图2。在混合对小鸡：马戏团aeruginosus x长
spilonotus在角32天，安加尔斯克，东西伯利亚，俄罗斯，16
1999年7月。皇冠显示较少黑暗比普通的条纹
羽毛的长spilonotus小鸡。 Tkis是诊断混合
父母子女关系。照片由本人。费费洛夫。
@ 2001'英国鸟类联盟，宜必思，143，587-592
590静脉注射费费洛夫
图3。在角鸡的鹞30天，伊尔库茨克，
东西伯利亚。俄罗斯。 1999年7月17日。它的羽色是
spilonotuschicks内的变异通常的限制，但稍微
接近'黑'色类型。摄影：一，费费洛夫。
'天空上升的'存在。有时第一phasr
顺利的secoiid，或'天空的变化，不断上升的'是
缺席。同样的显示屏也观察到
ncsting，但thcn他们大多都有一个地区性的功能。
一位女大约能参与男性飞行
展示技术！。男性长mwginosus显示圣公会舞只
（痉挛钛西蒙斯1 YSO）。一些消息来源（Dement'ev
1951年，科列洛夫1962）描述其天空显示舞
结尾的潜水和一降飞行要求。字母a
显示男性长育种近aeruginosus
安加尔斯克相似，长spilonotus，没有一
突然切换水平阶段的
降Ø我我吨'。
嵌套
荤jpilonotzis occiipip \为C uerugz相同的栖息地
nosus的pool5组成的'ibetlands在一个开放的
景观丰富紧急1 egetation（especiallk
里德，Reedmace ETI的1 111高数小
哺乳动物和鸟类8 \百分之巢是建立
Ø \呃11提尔Ninctv音响\百分之巢位置在色楞格河
dclta分布在Phraginztes芦苇丛，很少
上午十\吨\'在othcr高草密建（Fefelox欧文
1996）。在伊尔库茨克附近，其中里德缺席，沼泽鹞
经常窝阔叶Narrov叶
Reedmace。
虽然一夫多妻是常见的长在aeruginosus
欧洲的一些地区（席佩尔1979年，威特科斯基
1989年），并没有发现在C spilonotus。产卵
通常发生在5月10日在色楞格河三角洲
6月10日，在伊尔库茨克附近它开始前一个星期左右。字母a
完成离合器包含3-7蛋，通常是4-5
（费费洛夫1996j，这是奠定了2-3天外。的
平均离合器在色楞格河三角洲的大小各不相同4.3
5.3年之间，根据粮食供应（费费洛夫
1996）。这是接近长aeruginosus在欧洲
不同在法国平均窝卵数各地区
从四月1日至4月6日（Thiollay 1970年，Bavoux等。1989年），在
捷克共和国是4.77（Mattas 1988年）和
欧洲作为一个整体是4.0-4.4（Makatsch 1974）。
鸡蛋和身材
C的spiloriotus平均卵量测的
色楞格河三角洲是50.64 0.23（范围43.2-56.9）×
三十八点七一吨0.20（范围34.6-42.5）毫米，ñ = 91（平均吨
资深大律师）（费费洛夫1996年）。这些类似的鸡蛋
同样的形式从中国：49.5（45-54）× 38.6
（35.6-42.1）毫米（长和江1988j。C的鸡蛋
从northlvestern俄罗斯aeruginosus类似
那些长spilonotus（50.23（48.2-52.6）× 38.83
（37.2-40.1）毫米，周长= 26）（Il'inski等。1990年），以及
以作为一个整体（49.51 x三十八点五零毫米，ñ欧洲蛋
= 44 1焦耳（Makatsch 1974）。但是，它们更大的
斯堪的纳维亚和欧洲南部和较小
中欧（Makatsch 1974）。
C的spiloiiotus车身尺寸稍大于
那些中部和东部欧洲人群
长aeruginosus，但它们是相似的，或变小了
比那些在斯堪的纳维亚半岛和法国（Dement'ev
1951年，韦克1980年，Kavoux等。 1988年，这项研究）。
没有显着差异，在鸡蛋的证据
这些表格或身体大小。
繁殖成功
对C的平均成功巢窝雏数〜pilorio
在色楞格河drlta范围吨苞从八月一日至八月二号
之间的战争tledghngs（费费洛夫1996）meail
对所有筑巢大约1 8每双鸡，这是显着
在C比少aerugzizosus的河畔\ ival的
雏鸟取决于食物的供应
色楞格河三角洲，雏鸟mortalit！上升至31'％）和
繁殖成功（幼鸟/ eggj不同27日至
43％，这也是高于C为低nerugziiosus
2001年英国鸟类联盟/ B / S的143，587-592
选育的东部和西部沼泽鹞591
相比之下，繁殖成功的69-70％
Barabinskaya草原（科舍廖夫1983年），58％的普斯科夫
地区（Il'inski等。19903，2556在夏朗德省％
海事（法国）（Bavoux等。1989年）和41％
卡马格（法国）（Thiollay 1970）。
食物
一般来说，沼泽鹞是杂食性在大多数
地区（格雷贝尔德尔奥约等人。1994年），和C spilonotus没有
例外。小型啮齿动物（主要是田鼠属。）
形成的色楞格河三角洲的主要食品，马隆了
约80％的按编号饮食;鸟不能超过
20％的按编号饮食（费费洛夫1996年）。男性长
spilonotus喜欢小猎物采取若干lulometres
从巢，而女性会更接近的动物
对巢一样，其他鹞女性（席佩尔
1973）。然而，总质量的鸟类陷入更
比哺乳动物，由于禽流感的主要猎物
采取的是鸭和涉禽。在色楞格河三角洲，长spilonotus
看来要当鼠雀形目幼鸟
数字低。
讨论I0信噪比：比较与
系统在我吴长胜
长aeruginosus和C. spilonotus非常相似的大小，
筑巢生物学和饮食习惯，但他们中有很大的不同
羽毛着色。不过，这并不妨碍
他们的配对。杂交，与生产
对杂交种可能肥沃，确认在一个小
西伯利亚南部地区。养殖范围的大部分
填充表型'纯粹'鸟。最
可能的原因限制了混合区是最近的风景
历史在这些形式的接触面积，
因为针叶林和高山位置
南西伯利亚中部和西部蒙古提供
沼泽鹞的机会很少。斯坦波尼扬
（1990）认为，长期隔离嵌套
这些地区之前，他们最近的二次形式
联系方式。这种隔离期间可能出现
欧亚的更新世冰川阶段。
长spilonotus是，在许多方面，中间
与长aeruginosus和C. approximans。它具有
光臀（费费洛夫1996年）和发现下身
长approximans，但强烈的两性异形
在C aeruginosus。这二型是不存在在C
approximans（韦克1980）。该显示器的行为和
交配长spilonotus制度更类似于
一夫一妻制（和单态）和长ranivorus
'偶尔一夫多妻'长approximans比为C
aeruginosus。虽然女性长着色
spilonotus是介于长aeruginosus和
长approximans，即男性喜欢看
马达加斯加马什鹞（包括留尼汪岛和
马达加斯加形式长maillandi maillardi维氏
1862年，和C. maillardi macrosceles牛顿1863）
其射程，以及认为（3。ranivorus，很远
来自亚洲和大洋洲。这是非常有趣的
女长aeruginosus与部分带尾巴
羽毛有时记录在欧洲。
这些都是非常相似，长spilonotus（比曼＆
马奇1998年）。目前还不清楚他们是否长spilonotus，
或混合型，或者有悖常理的个人。少年长
spilonotus（见图。3）相似的长aeruginosus者
（费费洛夫1996年），只是spilonotus有更多的光线
羽毛对机翼，背部和臀部上侧;
有时完全黑暗的小鸡都记录在两个
形式。
毫无疑问，东马师必须鹞
位于相同的其他沼泽分类级别
鹞，构成西方supcrspecies，
澳大利亚，ReunionlMadagascan和非洲的形式，
的建议，在最近的作品（Arnadon＆红牛1988年，德尔
奥约等人。 1994）。可这一水平应当是一个'全'
物种？根据生物物种概念，至少，
我们应该考虑长aeruginosus和C. spilonotus
只需要亚种分化明显，因为
其杂交和事实，他们似乎
除了地理有任何障碍之一。
但是，这个问题的答案是最有可能
结果从分子系统学，比较研究
生态，动物行为学。
我很感谢很多同事谁帮助了我
编写本文件，即：医生V.A.波德科维罗夫，博士
1.1。 Tupitsin，V.E. Zhouravlyov和A.V. Shinkaryenko我们
曾在色楞格河三角洲了许多年，酋长的
命名aliove博士P.S.鸟类集合Tomkovich，
1.1。 Sirokhin，博士Ju.N. Nasarov'，和医生Ju.V. Bogorodsky博士
河西蒙斯博士W.J.A.席佩尔，克斯纳Cowing和人员
山猫Edicions（巴塞罗那），大肠杆菌;。大肠杆菌Curpian，请谁
我取得了一些论文和书籍。医生生态学家戈斯勒尔，编辑
在宜必思，和匿名裁判帮我一个文本
翻译成英语。还我应该特别感谢教授
O.M. Kozhova +，在伊尔库茨克的负责人生物研究所
大学在1989至1999年，为支持这项研究。
DISTRIBUTION OF MARSH HARRIERS IN
CENTRAL SIBERIA
The breeding range of C. aeruginosus extends east to
Lake Raikal, while that of the C. spilonotus extends
west to the Yenisey River and northwestern Mongolia
(Stepanyan 1990). This differs from the description
given by del Hoyo et al. (1 994), where these forms are
shown as allopatric. Marsh harriers are unevenly distributed
in the upper Yenisey valley (Kokhanovski
1991) and in the area between Yenisey and Baikal,
where they are more scarce, because of the shortage
here of the most productive wetlands, on which these
birds depend.
The Irkutsk region is populated by birds of both
forms. In its centre, C. aeruginosux is more frequent
(Mel'nikov 1999, this study), although these data are
insufficiently detailed; in the south and east (e.g. near
Irkutsk) C. spilonotus predominates, Some local sites of
C. aeruginosus in the Angara River valley (Skalon
1934) were flooded after the creation of the Bratsk
Reservoir in the 1960s. On the western shore of Baikal,
spilonotus is dominant (Fefelov 1996); in the Selenga
delta all birds of 132 observed nesting pairs were C.
spilonotus, and on only two occasions (1987 and 1990)
was a single adult male C. aeruginosus seen here
(Fig.1). There are no records of C. aemginosus in the
eastern Transbaikal or further eastwards.
HYBRIDIZATION OF EASTERN AND
WESTERN MARSH HARRIERS
The areas between Yenisey and Baikal, and northwestern
Mongolia are the zones of intergradation between
C. aeruginosus and C. spilonotus, which are populated
by phenotypically 'pure' birds of both forms as well as
Figure 1. Distribution of Circus aeruginosus and C. spilonotus in the south of Eastern Siberia. \\\, range of aeruginosus only; ///, range
of spilonotus only: cross-hatching, area inhabited by aeruginosus and spilonotus; ?, insufficient data; x, points where breeding of aeruginosus
was confirmed formerly or in the present time; n. points where breeding of spilonofus was confirmed; W , points of confirmed
inter-breeding of aeruginosus and spilonotus; -. -, state border of Russia; -. . . -, borders of Russian administrative regions.
0 2001 British Ornithologists' Union, Ibis, 143, 587-592
Breeding of Eastern and Western Marsh Harriers 589
by birds having intermediate plumage (Dement’ev
195 1, Stepanyan 1990). However, there is little information
on this. Both races show considerable
individual and age variation, which creates major difficulties
for field identification of possible hybrids.
From the classic descriptions of coloration of these
forms (Dement’ev 195 1, Stepanyan 1990, Weick
1980), and from my own observation, the most distinctive
field characters of adult c. aeruginosus are
dominance of brown on the belly and vent in males,
and lack of banding on the primaries, secondaries and
tail in females. For C. spilonotus, white predominates
on the belly and vent in males, while in females the
wing and tail feathers are banded and the rump is
whitish or pale. Another published character, that of
blaclush or brownish upperparts in males, is variable
and unclear. If we take these characters as the basis for
identification, no specimens are found that exhibit features
of both C. aeruginosus and C. spilonotus in the
collections named above. There is only one likely
hybrid adult male in the collection of Moscow
University, taken by N.M. Przevalsh on the Lobnor
Lake in Central Asia (undated), which is generally similar
to C. aeruginosus, but has the white vent and lower
belly of spilonotus (Fefelov 1999). In addition, adult
males with plumage intermediate between C. aeruginosus
and C. spilonotus were collected in northwestern
Mongolia (Khara-Usu Lake) and in western China
(Yarkend, Mazar-Tag) (Stegmann 1937). In general,
there are very few collected specimens from areas
where intergradation is likely to occur.
A ‘spotted’ type of male C. spilonotus that accounts
for 5-10% of nesting males in the Selenga delta
(Fefelov 1996) requires further research, as it is not
clear whether this plumage represents individual variation,
or is specific to 2- and 3-year-old birds. They
cannot be hybrids, however, because they are recorded
in different regions, including the eastern Transbaikal,
the Far East (in collections mentioned above) and in
Japan (Massey et al. 1990).
The natural shortage of good nesting sites and
human destruction of sites during the 20th century in
the zone of sympatry have reduced the likelihood of
inter-breeding between C. aeruginosus and C. spilono-
An adult male of apparently hybrid origin nested
near Irkutsk in 1993-95; this was probably the same
bird each year [Fefelov 19991. On the colour of the
tus.
spilonotus in all these years. At the same time from one
to two pairs of typical C. spilonctus nested here.
A nesting male C. aeruginosus was recorded near
Angarsk in 1995, 1998, 1999 arid 2000, paired with a
female C. spilonotus. It is uncertain whether these were
the same birds each year because they showed no individually
recognizable features iri their plumage. Their
nests were found in 1999 and 2000. Four chicks out of
five hatching eggs fledged successfully in 1999 (Fefelov
1999). Five chicks (two males and three females)
fledged in 2000. The plumage of hybrid fledglings
(Fig. 2) was very similar to those of C. spilonotus
(Fig. 3). The hybrid chicks have less dark streaking of
the crown than the majority of (3. spilonotus. However,
they cannot be discriminated in the field. In 1996 and
1997, a single pair of C. spilonotus nested in this area.
Thirty nesting cases of marsh harriers have been
recorded near Irkutsk and Angarsk during c. 20 years
(Ryabtsev & Fefelov 1997, this study). In 23 of them
both the parents were C. spilonotus, in four the male
was C. aeruginosus and in threc the male had hybrid
characteristics.
BREEDING ECOLOGY
Display
In spring, the courtship flight of C. spilonotus is either
very similar to that of the African Marsh Harrier C.
ranivorus Daudin 1800, and the Pacific (Australasian)
Marsh Harrier C. approximans l’eale 1848, (Simmons
1991), or it is intermediate between their display and
that of C. aeruginosus. In male and female C. spilonotus,
horizontal ‘sky-dancing’ and descending
upperparts, it could be either c, amginosus or C, spilonotus’
Its breast was coloured as in c’ spilonotusl and
the lower the vent and as in c. am&-
nosus. Its femalels?) and fledglings were coloured as C.
Figure 2. The chick of a mixed pair: Circus aeruginosus x C.
spilonotus at c. 32 days, Angarsk, Eastern Siberia, Russia, 16
July 1999. The crown shows less dark streaking than the usual
plumage of C. spilonotus chicks. Tkis is diagnostic of mixed
parentage. Photo by I . Fefelov.
@ 2001 British Ornithologists’ Union, Ibis, 143, 587-592
590 I.V. Fefelov
Figure 3. The chick of Circus spilonotus at c. 30 days, Irkutsk,
Eastern Siberia. Russia. 17 July 1999. Its plumage coloration is
within usual limits of variation of spilonotuschicks, but it is slightly
closer to 'dark' colour type. Photo by I. Fefelov.
'sky-spiralling' are present. Sometimes the first phasr
changes smoothly to the secoiid, or 'sky-spiralling' is
absent. The same displays are also observed during
ncsting, but thcn they mostly have a territorial function.
A female flying around can participate in a male
displa!. Male C. mwginosus show skh -dancing only
(Cramp ti Simmons 1 YSO). Some sources (Dement'ev
1951, Korelov 1962) describe its display as sky-dancing
ending with diving and calling in a descending flight. A
display of the male C. aeruginosus breeding near
Angarsk was similar to that of C. spilonotus without a
sudden switch from the horizontal phase to the
descending o i i t ' .
Nesting
C jpilonotzis occiipip\ the Same habitat as C uerugz
nosus consisting of pool5 and 'ibetlands In an open
landscape rich in emergent 1 egetation (especiallk
Reed, Reedmace eti 1 111th high numbers of small
mammals and birds Eight\ percent of nests are built
o\ er 11 ater Ninctv-fi\ e percent of nests in the Selenga
dclta are located in reedbeds of Phraginztes, rarely the
nr\t\ 'irv built in othcr dense tall grasses (Fefelox
1996). Near Irkutsk, where Reed is absent, marsh harriers
often nest in Broad-leaved and Narrov-leaved
Reedmace.
While polygyny is common in C. aeruginosus in
some areas of Europe (Schipper 1979, Witkowski
1989), it has not been found in C. spilonotus. Egg-laying
usually occurs in the Selenga delta from 10 May to
10 June, near Irkutsk it begins about a week earlier. A
complete clutch contains 3-7 eggs, most often 4-5
(Fefelov 1996j, which are laid 2-3 days apart. The
mean clutch-size in the Selenga delta varies from 4.3 to
5.3 between years, depending on food supply (Fefelov
1996). This is close to C. aeruginosus in Europe where
in various regions of France mean clutch size varies
from 4.1 to 4.6 (Thiollay 1970, Bavoux et al. 1989), in
the Czech Republic it is 4.77 (Mattas 1988) and in
Europe as a whole it is 4.0-4.4 (Makatsch 1974).
Egg and body sizes
Average egg measurements of C. spiloriotus from the
Selenga delta were 50.64 0.23 (range 43.2-56.9) x
38.71 t 0.20 (range 34.6-42.5) mm, n = 91 (mean t
sc) (Fefelov 1996). These are similar to eggs from the
same form from China: 49.5 (45-54) x 38.6
(35.6-42.1) mm (Chang & Jiang 1988j. Eggs of C.
aeruginosus from northlvestern Russia are similar to
those of C. spilonotus (50.23 (48.2-52.6) x 38.83
(37.2-40.1 ) mm, pi = 26) (Il'inski et al. 1990) as well
as to European eggs as a whole (49.51 x 38.50 mm, n
= 44 1 j (Makatsch 1974). However, they are larger in
Scandinavia and southern Europe and smaller in
Central Europe (Makatsch 1974).
Body sizes of C. spiloiiotus are a little larger than
those of central and eastern European populations of
C. aeruginosus, but they are similar to, or a little smaller
than, those in Scandinavia and France (Dement'ev
1951, Weick 1980, Kavoux et al. 1988, this study).
There is no evidence for significant differences in egg
or body sizes of these forms.
Reproductive success
The mean brood size of Successful nests of C ~pilorio
t u c in the Selenga drlta ranges from 1 8 to 2 8
tledghngs between wars (Fefelov 1996) The meail
over all nests is about 1 8 chicks per pair This is significantly
less than in C aerugzizosus The sur\ival of
nestlings depends on the availability of food In the
Selenga delta, nestling mortalit! was up to 31'%), and
breeding success (fledglings/eggj varied from 27 to
43% This i s also less than that of C nerugziiosus For
2001 British Ornithologists Union /b/s 143, 587-592
Breeding of Eastern and Western Marsh Harriers 591
comparison, breeding success is 69-70% in
Barabinskaya steppe (Koshelev 1983), 58% in Pskov
Region (Il’inski et al. 19903, 2556% in Charente-
Maritime (France) (Bavoux et al. 1989) and 41% in
Camargue (France) (Thiollay 1970).
Food
In general, marsh harriers are polyphagous in most
areas (del Hoyo et al. 1994), and C. spilonotus is no
exception. Small rodents (mostly voles Microtus spp.)
form the main food in the Selenga delta, malung up
about 80% of the diet by number; birds never exceed
20% of the diet by number (Fefelov 1996). Male C.
spilonotus prefer small prey taken several lulometres
from the nest, while females take larger animals nearer
to the nest, as do females of other harriers (Schipper
1973). However, the total mass of birds caught is more
than that of mammals because the main avian prey
taken is ducks and waders. In the Selenga delta, C. spilonotus
appear to take passerine fledglings when rodent
numbers are low.
DISCUSS I0 N : COMPARISON AND
SYSTEM AT I C S
C. aeruginosus and C. spilonotus are very similar in size,
nesting biology and diet, but they differ greatly in
plumage coloration. Nevertheless, this does not prevent
their pairing. Interbreeding, with the production
of hybrids which may be fertile, is confirmed in a small
area of southern Siberia. Most of the breeding range is
populated by phenotypically ‘pure’ birds. The most
likely cause limiting the hybrid zone is the recent landscape
history in the area of contact of these forms,
because the location of taiga forests and mountains in
southern central Siberia and western Mongolia offers
very few opportunities for marsh harriers. Stepanyan
(1990) considers that a prolonged isolation of nesting
areas of these forms preceded their recent secondary
contact. This isolation probably appeared during the
Eurasian glacial phases of the Pleistocene.
C. spilonotus is, in many respects, intermediate
between C. aeruginosus and C. approximans. It has the
light rump (Fefelov 1996) and spotted underparts of
C. approximans, but it is strongly sexually dimorphic as
in C. aeruginosus. This dimorphism is not present in C.
approximans (Weick 1980). The display behaviour and
mating system of C. spilonotus are more similar to the
monogamous (and monomorphic) C. ranivorus and
‘occasionally polygynous’ C. approximans than to C.
aeruginosus. Although the coloration of the female C.
spilonotus is intermediate between C. aeruginosus and
C. approximans, that of the male looks like the
Madagascar Marsh Harrier (including Reunion and
Madagascar forms C. maillandi maillardi Verreaux
1862, and C. maillardi macrosceles Newton 1863)
whose range, as well as that of (3. ranivorus, is very far
from Asia or Australasia. It is very interesting that
female C. aeruginosus with partially banded tail and
wing feathers are sometimes recorded in Europe.
These are very similar to C. spilonotus (Beaman &
Madge 1998). It is unknown whether they are C. spilonotus,
or hybrids, or aberrant individuals. Juvenile C.
spilonotus (see Fig. 3) resemble -those of C. aeruginosus
(Fefelov 1996) except that spilonotus have more light
feathers on the upper side of the wing, back and rump;
sometimes wholly dark chicks are recorded in both the
forms.
Undoubtedly the Eastern Marsh Harrier must be
located at the same taxonomic level as other marsh
harriers, which constitute a supcrspecies with Western,
Australasian, ReunionlMadagascan and African forms,
as suggested in recent works (Arnadon & Bull 1988, del
Hoyo et al. 1994). Can this level be that of a ‘full’
species? Under the Biological Species Concept at least,
we should consider C. aeruginosus and C. spilonotus
only to be significantly diverged subspecies because of
their hybridization and the fact that they appear to
have no barriers except the geographical one.
However, an answer to this question is most likely to
result from the study of molecular systematics, comparative
ecology and ethology.
I am grateful to many colleagues who helped me in the
preparation of this paper, namely: Dr V.A. Podkovyrov, Dr
1.1. Tupitsin, V.E. Zhouravlyov and A.V. Shinkaryenko as we
worked in the Selenga delta together for many years; chiefs of
ornithological collections named aliove Dr P.S. Tomkovich,
1.1. Sirokhin, Dr Ju.N. Nasarov’, and Dr Ju.V. Bogorodsky; Dr
R. Simmons, Dr W.J.A. Schipper, M.J. Cowing, and personnel
of Lynx Edicions (Barcelona), e.,;. E. Curpian, who kindly
obtained for me some papers and books. Dr A.G. Gosler, editor
of Ibis, and anonymous referees helped me with a text
translation to English. Also I owe gratitude especially to Prof.
O.M. Kozhova+, the head of Institute of Biology at Irkutsk
University during 1989-99, for the support of this research. |
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