2 research outputs found
Insight into structure: function relationships in a molecular spin-crossover crystal, from a related weakly cooperative compound
No full textThis is a repository copy of Insight into structure: function relationships in a molecular spin-crossover crystal, from a related weakly cooperative compound. White Rose Research Online URL for this paper: http://eprints.whiterose.ac.uk/83008/ Version: Accepted Version Article: Elhaïk, J, Kilner, C and Halcrow, MA (2014) Insight into structure: function relationships in a molecular spin-crossover crystal, from a related weakly cooperative compound. European Journal of Inorganic Chemistry, 2014 (26). 4250 -4253. ISSN 14344250 -4253. ISSN -1948 https://doi.org/10.1002/ejic.201402623 [email protected] https://eprints.whiterose.ac.uk/ Reuse Unless indicated otherwise, fulltext items are protected by copyright with all rights reserved. The copyright exception in section 29 of the Copyright, Designs and Patents Act 1988 allows the making of a single copy solely for the purpose of non-commercial research or private study within the limits of fair dealing. The publisher or other rights-holder may allow further reproduction and re-use of this version -refer to the White Rose Research Online record for this item. Where records identify the publisher as the copyright holder, users can verify any specific terms of use on the publisher's website. Takedown If you consider content in White Rose Research Online to be in breach of UK law, please notify us by emailing [email protected] including the URL of the record and the reason for the withdrawal request. Insight into Compound Jérôme Elhaïk, [a] Colin A. Kilner, [a] and Malcolm A. Halcrow* [a] Abstract: The ClO4 − salt of [FeL2] 2+ (L = 2,6-bis(3-methylpyrazol-1-yl)pyridine) undergoes very gradual thermal spin-crossover centered just below room temperature. In contrast, the BF4 − salt of the same complex exhibits an abrupt and structured spin-transition at lower temperature, with a complicated structural chemistry. The difference can be attributed to a much larger change in molecular structure between the spin states of the complex in the more cooperative BF4 − salt, leading to an increased kinetic barrier for their interconversion. Consistent with that suggestion, the high-spin and low-spin structures of weakly cooperative [FeL2][ClO4]2 are almost superimposable. The continuing interest in thermally and optically switchable spin-crossover (SCO) materials [9] Its thermal spin-transition takes place in two steps, via a re-entrant symmetry-breaking transition to an intermediate crystal phase, with a tripled unit cell containing a mixture of high-spin and low-spin sites. The first of these steps occurs abruptly with hysteresis, but at a temperature that varies according to the water content of the sample (x). In contrast the second step is kinetically slow, and is only achieved when the sample is poised at 100 K for 1.5 hrs. [10] Its excited spin-state trapping (LIESST [11] ) behavior is also unique, in that its thermodynamic high low spin transition and kinetically controlled high low spin-state relaxation exhibit different profiles and are effectively decoupled from each other. [12] Although unexceptional in itself, 1[ClO4]2 provides useful insight into the structural origin of the unusual behavior of the BF4 − salt by providing a rare comparison between strongly and weakly cooperative spin-crossover materials based on the same complex molecule. At 300 K, MT for 1[ClO4]2 is 2.4 cm 3 mol -1 K, lower than expected for a high-spin iron(II) complex with this ligand type (3.4-3.6 cm 3 mol -1 K)
A guide to nestling development and aging in altricial passerines
No full textThis is a guide to altricial passerines. It has an explanation of variables used for aging birds and an explanation of methodology employed, and provides information obtained while recording the early aging of the Dusky Flycatcher, the Carolina Wren, the Wrentit, Sprague’s Pipit, the Song Sparrow, the Chestnut-collared Longspur and the American Goldfinch.U.S Fish & Wildlife Service
A Guide to Nestling
Development and Aging
in Altricial Passerines
Biological Technical Publication
BTP-R6008-2007
U.S Fish & Wildlife Service
A Guide to Nestling
Development and Aging
in Altricial Passerines
Biological Technical Publication
BTP-R6008-2007
Dennis Jongsomjit1
Stephanie L. Jones2
Thomas Gardali1
Geoffrey R. Geupel1
Paula J. Gouse3
1 PRBO Conservation Science, Petaluma, CA
2 U.S. Fish and Wildlife Service, Region 6, Office of Migratory Birds,
Denver, CO
3 U.S. Fish and Wildlife Service, Bowdoin National Wildlife Refuge,
Malta, MT
Cover images: Top: Crissal Thrasher, Toxostoma crissale
Bottom: Brewer’s Sparrow, Spizella breweri
Photo credits: Top: Chris McCreedy/PRBO
Bottom: Colin Wooley/PRBO
ii A Guide to Nestling Development and Aging in Altricial Passerines
Author contact information:
Dennis Jongsomjit
PRBO Conservation Science
3820 Cypress Drive
Petaluma, CA 94954
707-781-2555
email:[email protected]
Stephanie L. Jones
U.S. Fish and Wildlife Service, Region 6
Nongame Migratory Birds
P.O. Box 25486 DFC
Denver, CO 80225
303-236-4409
Email: [email protected]
Thomas Gardali
PRBO Conservation Science
3820 Cypress Drive
Petaluma, CA 94954
415-868-0655 x 381
Email: [email protected]
Geoffrey R. Geupel
PRBO Conservation Science
3820 Cypress Drive
Petaluma, CA 94954
415-868-0655 x 301
Email: [email protected]
Paula J. Gouse
U.S. Fish and Wildlife Service
Bowdoin National Wildlife Refuge
194 Bowdoin Auto Tour Rd
Malta, MT 59538
406-654-2863
Email: [email protected]
For additional copies or
information contact:
U.S. Fish and Wildlife Service, Region 6
Nongame Migratory Bird Program
P.O. Box 25486 DFC
Denver, CO 80225-0486
Recommended citation:
Jongsomjit, D., S. L. Jones, T. Gardali, G. R.
Geupel, and P. J. Gouse. 2007. A guide to nestling
development and aging in altricial passerines. U.S.
Department of Interior, Fish and Wildlife Service,
Biological Technical Publication, FWS/BTP-R6008-
2007, Washington, D.C.
Table of Contents iii
Table of Contents
List of Figures. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iv
Acknowledgments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vi
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Nestling Growth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Analyzing Growth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Growth Patterns and Aging . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Aging Recommendations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Variables Used for Aging. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Individual feather tracts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Wing chord. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Weight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Tarsus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Primary and rectrices pin lengths. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Culmen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Eyes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Longest broken primary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Total body length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Gape and rictus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Physical and behavioral descriptors. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Objectives. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Using the Species Accounts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Measurements and Terminology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Feather definitions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Weight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Tarsus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Wing chord. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Primary and rectrix pin length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Culmen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Total length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Longest broken primary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Gape and rictus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Physical and behavioral descriptors. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Species Account Tables. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Species information. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Indicator table. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
General feather development. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Key visual indicators. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Photographs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Feather tract data. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Morphometric data. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
General description. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Species Accounts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Dusky Flycatcher . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Carolina Wren. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Wrentit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Sprague’s Pipit. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Song Sparrow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Chestnut-collared Longspur. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
American Goldfinch. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Literature Cited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Appendix A. Adult Morphometric Measurements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Appendix B. Data Collection Methods and Protocols. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
iv A Guide to Nestling Development and Aging in Altricial Passerines
List of Figures
Figure 1. Dorsal and ventral views of a nestling with the individual feather tracts marked and identified. . . . .5
Figure 2. Tarsus with measurement indicators at the tibiotarsus joint and distal end of the last leg scale . . 6
Figure 3. Wing chord as measured with a wing ruler, without flattening or pressing down on the wing .. . . . . 6
Figure 4. Pin lengths as measured from the point of emergence from the skin . . . . . . . . . . . . . . 6
Figure 5. The culmen as measured from nares to tip . . . . . . . . . . . . . . . . . . . . . . . . . 6
Figure 6. Total body length taken from the tip of the bill to the tail bud . . . . . . . . . . . . . . . . . 6
Figure 7. The exposed portion of the longest broken primary as measured with a ruler. . . . . . . . . . . . . . . . . . . .6
Figure 8. The gape as measured with calipers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
Figure 9. Examples of feather development and appearance using a Wrentit on Day 7 . . . . . . . . . . 7
Figure B-1. An example of systematic labeling of data sheets and photographs which uniquely identifies
each nestling and their age. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Acknowledgments v
Acknowledgments
The collection of data for this guide would not have
been possible without the help of many interns and
field biologists including Parvaneh C. Abbaspour,
Bethany S. Atchley, Amon J. Armstrong, Ross R.
Conover, Gerhard A. Epke, Tristan R. Gingerich,
Geetha M. Jayabose, Scott F. Jennings, Laura H.
Kaplan, Errin B. Kramer-Wilt, Ben G. Larson,
Blaine MacDonald, Emily Morrison, Henry
Ndithia, Kerry E. Neijstrom, Alexander Rosenthal,
Corrina Snow, Amanda B. Shults, and Dionne R.
Wrights. Many thanks to Suzanne Austin-Blythell,
Bruce Barbour, Thomas M. Haggerty, Steve N. G.
Howell, and Diana L. Humple for providing helpful
comments to earlier drafts of this guide. Fig. 1 is
adapted from Gill (1994) and Fig. 6 is adapted from
Baldwin et al. (1931). We thank Peter Pyle and
Steve N. G. Howell for graciously allowing us the
use of Figs. 2, 3, and 5 (Pyle 1997). Figures 4, 7, and
8 were drawn by Dennis Jongsomjit. We thank the
Point Reyes National Seashore for their continued
cooperation. Funding was provided by the U.S. Fish
and Wildlife Service Region 6 Nongame Migratory
Bird Program. This is PRBO contribution 1603.
Data Contributors
We especially thank the colleagues that contributed
nestling data to this publication: Ryan Burnett and
Vanessa Tissdale – PRBO Conservation Science,
Dusky Flycatcher (Empidonax oberholseri) and
Thomas M. Haggerty – Department of Biology,
University of North Alabama, Carolina Wren
(Thryothorus ludovicianus).
1
Nestling growth and development studies have
been a topic of interest for a greater part of the
last century (Sutton 1935, Walkinshaw 1948)
and continue to be of interest today. This is not
surprising since studies on nestling growth can
provide a wealth of biological information that
has larger implications for avian management
and conservation. Despite this history of studying
nestling development, basic information is still
limited or absent for many species. Many questions
remain unanswered, and contradictory conclusions
are often found in the literature (Starck and Ricklefs
1998a). Therefore, much information on aging and
development can still be gained from studying the
development patterns of similar species and from
comparative studies, across avian orders (Minea
et al. 1982, Saunders and Hansen 1989, Carsson
and Hörnfeldt 1993). Additionally, nestling growth
studies can yield insight into the effects of different
nesting strategies on productivity (O’Connor
1978), as well as the impacts of parental effort and
environmental variables on fitness (Ross 1980,
Ricklefs and Peters 1981, Magrath 1991). Since low
reproductive success may play a significant role in
the declines of many North American passerines
(Sherry and Holmes 1992, Ballard et al. 2003), a
better understanding of the factors that influence
reproductive success is a vital component of avian
conservation (Martin 1992). Data on nestling aging
can be used to improve nest survival estimates
(Dinsmore 2002, Nur et al. 2004), providing
information that can be used to more precisely age
nests (Pinkowski 1975, Podlesack and Blem 2002),
(Jones and Geupel 2007). Indeed, the relatively short
time period young spend developing in the nest is
a critical part of a bird’s life cycle and a nestling’s
developmental path can affect its survival to
independence, its survival as an adult, and its future
reproductive success.
Nestling Growth
Ornithologists categorize birds over an altricial to
precocial spectrum, based on differences in the rate
of growth and type of development young birds will
undertake (Starck and Ricklefs 1998b). Placement
into this spectrum depends on various broad
characteristics such as mobility, feeding behavior,
presence of down, and parental nest attendance (Gill
1994). Growth rates are highly variable within the
altricial to precocial spectrum, with developmental
periods varying as much as 30-fold (Ricklefs 1983).
Much of this variation in growth can be attributed, at
the phylogenetic level, to differences in body mass.
In general, altricial species can grow at 3-4 times the
rate of precocial species, and growth rates of birds
with similar mass can vary by as much as a factor of
10 (Ricklefs et al. 1998). In this guide, we focus on
altricial species.
Nestling growth variability has largely been studied
looking at effects on individual fitness of offspring
and parents (e.g., Murphy 1983, Magrath 1991,
Halupka 1998). Differences between populations can
manifest as morphological differences or differential
timing in the growth and maturation of certain body
components (Murphy 1983, Burns 1993). However,
the growth rate of a single species throughout its
entire range can sometimes vary little (King and
Hubbard 1981, Murphy 1983, Pereyra and Morton
2001).
Variability in nestling growth rates can be due to
many ecological factors, in conjunction with specific
species life history strategies; some developmental
processes might be linked and are also independent
of the nutritional state of a nestling (Ricklefs
1968a). Some factors associated with species specific
growth rates and patterns include nest location,
synchronicity of hatching, and brood size (Murphy
1983). Ecological factors that influence nestling
growth are generally related to limitations in food
availability (Ricklefs 1993), including weather
(Petersen et al. 1986), habitat differences and quality
(Ross 1980, Dawson and Bidwell 2005), parasites
(Burhans et al. 2000), competition between nest
mates (Werschkul and Jackson 1979, Ricklefs 1982),
and parental abilities (Briskie 1995). Additionally,
higher nest-predation rates may favor higher
nestling growth rates (Lack 1968, Remes and Martin
2002, but see Ricklefs 1969).
At the physiological level, an important factor
thought to limit growth is “tissue level constraint”,
where nestlings are growing at a maximum rate
allowable by the tradeoff existing between resources
available for growth and mature tissue function
(Ricklefs et al. 1998). Once certain types of cells
differentiate into mature functioning tissue, they
no longer continue to grow (O’Connor 1984). After
a period of below normal growth, a nestling would
need to increase its growth rate in order to “catch
up” to its normal developmental timing. However,
such compensatory growth has not been shown to
occur in altricial birds (Schew and Ricklefs 1998,
Lepczyk and Karasov 2000; but see Remes and
Martin 2002, Bize et al. 2006). In one study, addition
of body mass and growth rates in overfed young in
the laboratory did not differ from that of wild young
Introduction
Introduction
2 A Guide to Nestling Development and Aging in Altricial Passerines
(Konarzewski et al.1996). These results indicate that
young may be growing at the maximum rate allowed
by cell function and physiology.
Analyzing Growth
An important part of visualizing and analyzing
nestling growth is the use of fitted growth equations
(Ricklefs 1967, 1983). When fitted into a growth
equation, using non-linear regression, three
components of growth are provided: the rate,
magnitude, and form of growth. When graphed,
nestling growth is often shown to increase, reach
a peak, and finally level off in a sigmoidal shape.
These equations simplify and allow for comparative
analysis of growth between and within species.
Information on adult body weight and size are
also an important aspect of analyzing growth with
these equations (O’Connor 1984); adult body size
measurements are provided for the study species in
this guide (Appendix A). Alternatively, growth data
can also be used to build predictive models of age
(Holcomb and Twiest 1971, Hamel 1974).
Growth Patterns and Aging
Inherent species specific patterns of growth and
development can often be used to age nestlings
(Starck and Ricklefs 1998a). In nestling growth
patterns, each body component can begin growth
at a different point in time relative to other
components, resulting in a staggered growth
pattern. Specific patterns in this type of growth
are generally adaptations for nest survival. For
example, in some species, contour feathers tend to
rapidly grow and unsheathe before the remiges,
providing important insulation cover early in life,
when young cannot self-thermoregulate (Murphy
1981). In another example, growth of the tarsus or
gape, important for food acquisition, may proceed
rapidly during the early nestling stage (O’Connor
1984). Besides growth, developmental events
(e.g., pin-feather eruption patterns, eye opening,
and behavior) can be age specific and are readily
observed (Ricklefs 1966, Murphy 1981). Thus, using
a combination of several growth measurements
can provide reliable aging throughout the nestling
period (King and Hubbard 1981, Murphy 1981,
Haggerty 1994, Podlesak and Blem 2002).
Aging Recommendations
The type and number of measurements needed
for reliable aging may vary among species but
preliminary analysis of our data shows that wing
length, tarsus length, weight, and culmen provide
good predictive models of age. Since nestlings may
be growing at a maximized rate, age estimates can
be informed by considering the development of the
most advanced nestlings, and by using more than
one nestling. In nests parasitized by Brown-headed
Cowbird (Molothrus ater), where all the young
may be receiving less food than normal, aging host
young may or may not be reliable (Kilner et al. 2004,
Weatherhead 1989). Aging of the Brown-headed
Cowbird young may be possible, depending on the
host species (Scott 1979, Kilpatric 2002).
With any aging technique, it is important to be
aware that deviations from normal growth and
development can occur, preventing accurate
predic
