Commentary: A Recommendation for Standardized Age-Class

Plumage Terminology for Raptors

Author(s): William S. ClarkPeter Pyle
Source: Journal of Raptor Research, 49(4):513-517.
Published By: The Raptor Research Foundation
DOI: http://dx.doi.org/10.3356/rapt-49-04-513-517.1
URL: http://www.bioone.org/doi/full/10.3356/rapt-49-04-513-517.1

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 SHORT COMMUNICATIONS

J. Raptor Res. 49(4):513–517

E 2015 The Raptor Research Foundation, Inc.

COMMENTARY: A RECOMMENDATION FOR STANDARDIZED AGE-CLASS PLUMAGE TERMINOLOGY

FOR RAPTORS

WILLIAM S. CLARK1
2301 South Whitehouse Circle, Harlingen, TX 78550 U.S.A.

PETER PYLE
The Institute for Bird Populations, P.O. Box 1346, Point Reyes Station, CA 94956 U.S.A.

KEY WORDS: Age terminology; immature; juvenile; molt cycle; regions or in the southern hemisphere, where hatching
plumage; subadult. and fledging can span 1 January in a given species.
We recommend that standard age-class terminology be Additional sets of age terms that apply to raptors are
used in publications that involve raptor plumages. We pro- available for use in lieu of “subadult” and “immature”
pose using standardized terminology developed by Hum- and instead of calendar-based systems. All such sets have
phrey and Parkes (1959) and refined by Howell et al. clearly and unambiguously defined terms that can be used
(2003) for all raptors, and we propose age-coding terms for the various definitions above. We propose that the
recommended by Wolfe et al. (2010) and Johnson et al. Humphrey and Parkes (1959) molt and plumage terminol-
(2011), as described herein. ogy, as modified by Howell et al. (2003; the “H-P-H” ter-
The age terms “subadult” (or “sub-adult”) and “im- minology) be used to indicate age classes in raptors rather
mature” are used frequently in raptor articles, raptor than vague and/or calendar-based terms. We further pro-
books, and raptor field guides, and often in the raptor pose that for instances when the age of the raptor cannot
sections of bird field guides. However, the uses and defini- be determined, that the term “pre-definitive” be used for
tions of these terms have not been consistent among these all non-adult plumages.
publications. “Subadult” has been used to refer to birds in H-P-H Terminology Applied to Raptors. H-P-H terminol-
at least four different age/plumage classes: (1) the first ogy takes a “cycle-based” approach, in which a molt cycle
adult plumage when juvenile feathers are present (e.g., or plumage cycle is defined as the period between the
Clark and Wheeler 2001); (2) the annual plumage preced- beginning of one prebasic molt to the beginning of
ing adult plumage (e.g., Clark 1999); (3) all plumages the next prebasic molt (usually, but not always, a year).
between juvenile and adult (e.g., Liguori 2011, Wheeler The growth of first pennaceous feathers in the nest is con-
2003); and (4) all non-adult plumages (e.g., Newton sidered the first prebasic molt, and the period between this
1979). The term “immature” has been applied to raptors and the second prebasic molt (at about a year of age) is
in at least two ways: (1) for all non-adult plumages (e.g., considered the first molt cycle, followed by the second
Wheeler 2003), and (2) for any plumage between juvenile molt cycle, and so on.
and adult (e.g., Friedmann 1950; see also Clark 1999). Older raptors, including most breeding adults, undergo
However, the second definition has been used much less a single prebasic molt per cycle (typically a year), unlike
frequently. “Immature” means literally “not adult.” some bird species that have inserted prealternate molts
Most books and articles on ageing raptors for banding and two or more plumages per adult cycle (e.g., one plum-
or ringing use calendar-based ageing terms (e.g., Baker age during the breeding season and another plumage
1993, Bloom and Clark 2001, and Pyle 2008 in part). Un- outside the breeding season, such as in the American
der such terminology, age classes advance on 1 January Goldfinch [Spinus tristis]). In the first molt cycle, some
each year, despite no concurrent change in age-determi- raptors, such as New World vultures and perhaps some
nation criteria. Such terminology is required for banding larger eagles and falcons, appear to lack an inserted molt
in North America (Canadian Wildlife Service and U.S. Fish in all cycles, resulting in the Simple Basic Strategy (SBS) as
and Wildlife Service 1991), and similar calendar-based age defined by Howell et al. (2003). However, most raptor
coding is used in Europe and elsewhere. But such age-class species appear to have a limited or partial inserted molt
terminology becomes much more confusing in equatorial during their first molt cycle (Pyle 2005a), as is found in
most birds, and now referred to as the preformative molt.
1 Email address: [email protected] Most raptors, therefore, follow the Complex Basic Strategy

513
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Table 1. Comparison of cycle-based age-classification system codes recommended for North American raptors and
equivalents using the calendar-based age-classification system, for the first, second, and definitive molt cycles. Calendar-
based age codes are currently recognized by the U.S. Bird Banding Laboratory (BBL) for north-temperate breeding birds.
The BBL (http://www.pwrc.usgs.gov/BBL/MANUAL/index.cfm) is currently adopting WRP classification codes as well.
Note that some species have identifiable fourth-cycle (4CB) and fifth-cycle (5CB) basic plumages as well. Table adapted
from that of Pyle et al. (2015).

MOLT-BASED AGE-CLASSIFICATION SYSTEM CALENDAR-BASED AGE-CLASSIFICATION SYSTEM

FCJ 1st molt cycle, juvenile plumage HY Hatching year
FPF 1st molt cycle, undergoing PF molt HY Hatching year
FCF 1st molt cycle, formative plumage HY/SY Hatching year/2nd year
SPB 2nd molt cycle, undergoing 2nd PB molt SY 2nd year
SCB 2nd molt cycle, basic plumage SY/TY 2nd year/3rd year
TPB 3rd molt cycle, undergoing 3rd PB molt TY 3rd year
DPB Definitive molt cycle, undergoing definitive prebasic molt AHY/ASY After hatch year/after 2nd year
DCB Definitive molt cycle, basic plumage AHY/ASY After hatch year/after 2nd year

(CBS) according to Howell et al. (2003). Note that the Fifth, Sixth, etc. Prebasic Molts); DPB (undergoing the
term CBS refers to the species, but not all individuals of Definitive Prebasic Molt); and DCB (Definitive Basic Plum-
the species undergo a preformative molt; in other words, age). Various codes for “unknown” plumages are also avail-
the molt can vary from absent to limited within a species able (see Johnson et al. 2011, Pyle et al. 2015). Although
(Pyle 2005a). these codes were derived primarily for tropical species in
In H-P-H terms, raptors during the first molt cycle can which calendar-based age-coding systems are not useable,
be in “Juvenile Plumage” throughout the first cycle in SBS the codes can equally be applied to temperate North
species and some individuals of CBS species, or “Juvenile American species and this coding system will be intro-
Plumage” followed by “Formative Plumage” in most indi- duced as an option by the Bird Banding Laboratory
viduals of CBS species. In all raptor species this is followed (BBL; J. Lutmerding pers. comm.; Table 1). The WRP
by “Basic II” or “Second Basic” plumage during the second codes can be used to compare molts and plumages of
molt cycle, “Basic III” or “Third Basic” plumage during the raptors with the Simple Basic Strategy to those with the
third molt cycle, and so on, until “Definitive Basic” plum- Complex Basic Strategy, and to define age-groups as based
age is attained, the plumage that remains essentially static on the molting strategies of each species (Fig. 1).
from year to year and is found in most breeding adults. Summary of Molts, Plumages, and Proposed Age Desig-
This can occur variously among raptor species during the nations and Codes. Juvenile plumage (FCJ). This is the
second cycle beginning at 1 yr old (e.g., kites, some other plumage acquired in the nest after the downy stages and
smaller Accipitrids, and falcons) or during the fifth or later is synonymous with “First Basic Plumage.” It is the first
cycle beginning at five or later years of age (e.g., in Bald plumage of a raptor and in some species can be worn for
Eagle [Haliaeetus leucocephalus], and California Condor almost a year, throughout the first cycle, until commence-
[Gymnogyps californianus]); it is not always attained during ment of the Second Prebasic Molt (SBS species and some
the same cycle in all individuals (Pyle 2008). Thus, knowl- individuals of CBS species). All North American raptors
edge of specific maturation rates are needed to under- have distinct juvenile plumages, although those of male
stand the minimum age for which Definitive Plumage and female American Kestrels (Falco sparverius) differ only
can be applied, and it is not a term that can be assumed slightly from later plumages of each sex.
to represent homology between species or paralogy within Formative plumage (FCF). This plumage results from a lim-
species (Howell and Pyle 2014). ited-to-incomplete Preformative Molt, inserted within the
For age-coding in the field or in databases, a system first cycle of raptors showing the CBS molt strategy. Birds
based on H-P-H terminology has been derived by Wolfe undergoing the Preformative Molt can be coded FPF. This
et al. 2010), the “WRP system” as modified by Johnson et plumage occurs in some to many individuals of most to all
al. (2011). See Pyle et al. (2015) for a thorough treatment North American raptors, between juvenile and second-
of its use. For diurnal raptors, applicable plumage and basic plumages (Pyle 2005a), but has not been observed
molt codes are: FCJ (First Cycle, Juvenile Plumage); FPF in New World vultures. In most species, formative plumage
(First Cycle, undergoing the Preformative Molt); FCF resembles juvenile plumage, but with a few newer feathers
(First Cycle, Formative Plumage); SPB (Second Cycle, un- in the upperparts and underparts, which may resemble
dergoing the Second Prebasic Molt); SCB, TCB, 4CB, 5CB, definitive plumage or be intermediate between juvenile
etc. (Second, Third, Fourth, Fifth, etc. Basic Plumages); and definitive plumage. Three North American species,
TPB, 4PB 5PB, 6PB, etc. (undergoing the Third, Fourth, American Kestrel, Mississippi Kite (Ictinia mississippiensis),
DECEMBER 2015 SHORT COMMUNICATIONS 515

Figure 1. Comparison of molt cycles and plumages of Simple Basic Strategy with Complex Basic Strategy using
WRP codes.

and White-tailed Kite (Elanus leucurus), have more exten- most kites, and Red-shouldered Hawk (Buteo lineatus), sec-
sive preformative molts that include most or all body feath- ond basic plumage corresponds to definitive basic plum-
ers and, in the White-tailed Kite, can include flight feath- age and most or all birds after the second prebasic molt
ers as well (Pyle 2005a, 2008). In these species, formative are coded DPB as opposed to SPB. Other species, includ-
body plumage approaches or resembles definitive plum- ing eagles, condors, caracaras, and larger Buteonines, can
age. have distinctive second-basic body plumages. In still other
Basic II or second basic plumage (SCB). This is the second species, including Accipiters, harriers, and most Buteos,
full plumage of a raptor, following the second prebasic body plumage may or may not resemble definitive-basic
molt, which is complete or nearly complete in North body plumage, but second-basic plumage can be identified
American species, usually at 10 to 15 mo of age. Individuals by retained juvenile outer primaries and/or secondaries in
undergoing the second prebasic molt can be coded SPB. some or all individuals (Clark 2001, Bloom and Clark 2001,
In some North American species, including most falcons, Clark and Bloom 2005, Pyle 2005b, 2006, 2008).
516 SHORT COMMUNICATIONS VOL. 49, NO. 4

Basic III or third basic plumage (TCB). This is the third full COMENTARIO: RECOMENDACIÓN PARA EL USO DE
plumage of a raptor, acquired by a complete or incomplete U N A T E R M I N O L O G I´ A E S T A N D A R I Z A D A D E
third prebasic molt at about 2 yr of age; birds actively un- PLUMAJES DE CLASES DE EDAD PARA RAPACES
dergoing this molt can be coded TPB. Third-basic body
plumage resembles definitive basic plumage in most North RESUMEN.—Los términos “subadulto” e “inmaduro” han
American species (for which TCB cannot be coded) but sido aplicados en forma vaga e inconsistente en la literatura
differ markedly in Bald Eagle and California Condor, sobre rapaces. Además, los sistemas de registro de envejeci-
miento basados en el calendario generan cambios en los
and subtly in a few other larger species. In the eagles,
códigos a pesar de que no ocurren cambios en la apariencia
condor, Osprey (Pandion haliaetus), White-tailed Hawk
exterior u otros criterios utilizados para determinar la edad, y
(Geranoaetus albicaudatus), and Rough-legged Hawk (Buteo
no pueden ser utilizados en las regiones ecuatoriales donde
lagopus), third-basic plumage in some or all individuals
las aves pueden reproducirse desde el uno de enero. Para
can be identified by retained juvenile outer primaries or
clarificar y añadir consistencia a la terminologı́a de muda y
medial secondaries along with two generations of basic
de plumaje en rapaces, presentamos un grupo estandarizado
feathers in staffelmauser (stepwise) patterns, as well as
de términos de clases de edad y de códigos para reemplazar
pre-definitive body plumages in some species (Bloom
los términos que no han sido definidos universalmente. Pro-
and Clark 2001, Clark 2004, Clark and Bloom 2005, Pyle
ponemos una terminologı́a basada en un ciclo y una codifi-
2005a, 2006, 2008).
cación que aplique una nomenclatura precisa sobre la muda
Basic IV, V, and VI, or fourth, fifth, and sixth basic plumages y el plumaje para aves rapaces de las que se conoce la edad, y
(4CB, 5CB, and 6CB). These plumages follow the fourth, presentamos una cronologı́a de mudas y plumajes, ası́ como
fifth, and sixth prebasic molts, which are incomplete to sus códigos, para su uso en rapaces de América del Norte. En
complete and occur at about 4, 5, and 6 yr old, respective- los casos en los que la edad real no pueda ser determinada,
ly. Birds undergoing these molts can be coded 4PB, 5PB, proponemos el uso del té rmino “pre-definitivo” (o
and 6PB. Among North American raptors, only Bald Eagle “inmaduro”) para rapaces que no son ni adultos ni juveniles.
and California Condor have distinct fourth-basic plu- [Traducción del equipo editorial]
mages, some individuals of these species have recognizable
fifth-basic plumages, and occasional individuals may show ACKNOWLEDGMENTS
recognizable sixth-basic plumages. In all other North We thank Lloyd Kiff and several anonymous reviewers
American species these plumages are not distinguishable for helpful comments on earlier versions of this commen-
from Definitive Basic Plumage. tary. This is contribution number 497 of The Institute for
Bird Populations.
Definitive basic plumage (DCB). This is the final mature
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