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Rodrigo Barban Zucoloto
Patricia Susana Amavet
Luciano Martins Verdade
Izeni Pires Farias Editors
Conservation
Genetics
of New World
Crocodilians
Conservation Genetics of New World Crocodilians
Rodrigo Barban Zucoloto
Patricia Susana Amavet
Luciano Martins Verdade • Izeni Pires Farias
Editors
Conservation Genetics of
New World Crocodilians
Editors
Rodrigo Barban Zucoloto Patricia Susana Amavet
Laboratório de Genética de Populações e Laboratorio de Genética, Departamento
Evolução Molecular (GENEV), de Ciencias Naturales, Facultad de
Instituto de Biologia, Humanidades y Ciencias, Universidad
Universidade Federal da Bahia, Nacional del Litoral, Consejo Nacional
Salvador, Bahia, Brasil de Investigaciones Científicas y
Técnicas (CONICET),
Luciano Martins Verdade Santa Fe, Argentina
Universidade de São Paulo
CENA/LE²Ave Izeni Pires Farias
Piracicaba, São Paulo, Brasil Laboratório de Evolução e Genética Animal
Departamento de Genética
Instituto de Ciências Biológicas,
Universidade Federal do Amazonas,
Manaus, Amazonas, Brasil
ISBN 978-3-030-56382-0 ISBN 978-3-030-56383-7 (eBook)

https://doi.org/10.1007/978-3-030-56383-7
© Springer Nature Switzerland AG 2021

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Foreword
Since decades, information on crocodilians in the Americas has been available. The
champions of crocodilian conservation were Eduard Mc Ilhenny in the USA, Miguel
Alvarez del Toro in México, Federico Medem in Colombia, and Angel Gallardo in
Argentina, just to mention a few, with obviously many unfair omissions. These
examples, in my opinion, somehow describe what it was like to do science at that
time in our region, when solitary naturalists with or without a budget invested their
best to search, investigate, and communicate their findings mainly on the natural
history of our crocodiles.
Not so long ago, the Group of Crocodile Specialists with its meetings around the
world, especially on this continent, with the USA, Venezuela, Ecuador, and
Argentina being the first to participate, launched a process of consolidation of a
network of American crocodile specialists. In this way, the first volume on the con-
servation and management of Latin American alligators and crocodiles was pro-
duced in 1995, and from there, an increasing number of publications referring to
crocodiles were developed in different peer-reviewed journals. These publications
were characterized by multi-thematic content but with single authorship in
most cases.
Right now, with a thematic specialization and globalized world, and with easy
and cheap communication between researchers, more and more research is being
generated by multiple authors, very often belonging not only to different institutions
but also to different continents, as can be seen in this book. In my opinion, this gen-
erates more and better information on the one hand, but also favors its dissemination
at the local, regional, and international levels.
We were used to seeing in the old days that the information in South America
referred mainly to individual descriptions about a nesting female, the size of an
individual, or the finding of a crocodile in a place that allowed to determine the
approximate geographic distribution of that species. Despite the fact that the situa-
tion in North America was different, the publications referred to only one or two
species, which was totally useless for a conservation strategy at the regional level.
Globalization has many pros and cons, but from a scientific point of view, bene-
fits are almost the rule. In this context, we are also involved in a kind of
v
vi Foreword
“continentalization” process, and this book is an excellent example of this. The

synergy between authors from all over the Americas (and also from other conti-
nents) is generating high-quality results in topics such as hybridization, genetic
diversity, genetics and evolution, multi-paternity, and molecular phylogenetics,
among other topics, allowing the joint use of the best available technology no matter
the boundaries between countries.
I used to say many years ago that conservation programs for crocodiles must be
developed on a regional basis, because the species has no nationality and distribu-
tion area has no political boundaries, well, it seems that now this is also valid for
science and scientists. I cannot wait to see how things evolve in the near future. In
the meantime, I’m sure you’ll enjoy this book very much.

Departamento de Ciencias Naturales, Facultad de Alejandro Larriera
Humanidades y Ciencias
Universidad Nacional del Litoral
Santa Fe, Argentina
Proyecto Yacaré, Laboratorio de Zoología Aplicada:
Anexo Vertebrados (FHUC-UNL/MAyCC)
Santa Fe, Argentina
Acknowledgments
We thank all contributors of this book for working together to organize these texts
carefully and sharing decades of research effort in conservation genetics studies on
crocodilians.
vii
Contents
1 Geographic Distribution, Habitat, Reproduction,

and Conservation Status of Crocodilians in the Americas�� 1
Francisco Villamarín, Armando H. Escobedo-Galván,
Pablo Siroski, and William E. Magnusson
2 Molecular Markers Applied to Conservation Genetics
of American Crocodilians�� 31
Rodrigo Barban Zucoloto, Izeni Pires Farias, and Patricia
Susana Amavet
3 Molecular Phylogenetics of the New-World Crocodylia�� 79
Llewellyn D. Densmore III and Tomas Hrbek
4 Biogeography and Comparative Phylogeography
of New-World Crocodylians�� 95
Fábio de Lima Muniz, Pedro Senna Bittencourt, Sandra Marcela
Hernández-Rangel, Igor Joventino Roberto, Izeni Pires Farias,
and Tomas Hrbek
5 Genetic Diversity of New World Crocodilians�� 123
Patricia Susana Amavet, Rodrigo Barban Zucoloto, Tomas Hrbek,
and Izeni Pires Farias
6 Crocodilians Are Promiscuous But Not to the Benefit of
Heterozygosity�� 153
Sally R. Isberg
7 Hybridization and Speciation Among New-World
Crocodilian Species�� 171
Gualberto Pacheco-Sierra and Patricia Susana Amavet
8 Crocodilian Genome Advances �� 185
Katherine Brittain, David A. Ray, and Jaime Gongora
ix
x Contents
9 How Genetic Tools Can Help Crocodilians’ Management

and Governance�� 203
Luciano M. Verdade, Carlos I. Piña, Melina Simoncini,
and Karina L. Silva-Brandão
10 Perspectives and Final Considerations About the Molecular
Ecology of New-World Crocodilians�� 215
Patricia Susana Amavet and Rodrigo Barban Zucoloto
Index�� 223
Contributors
Patricia Susana Amavet Laboratorio de Genética, Departamento de Ciencias

Naturales, Facultad de Humanidades y Ciencias, Universidad Nacional del Litoral,
Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET),
Santa Fe, Argentina
Pedro Senna Bittencourt Universidade Federal do Amazonas. Departamento de
Genética, Manaus, Amazonas, Brasil
Katherine Brittain Sydney School of Veterinary Science, Faculty of Science,
University of Sydney, Sydney, NSW, Australia
Fábio de Lima Muniz Departamento de Genética, Universidade Federal do
Amazonas, Manaus, Amazonas, Brasil
Llewellyn D. Densmore III Department of Biological Sciences, Texas Tech
University, Lubbock, TX, USA
Armando H. Escobedo-Galván Centro Universitario de la Costa, Universidad de
Guadalajara, Puerto Vallarta, Jalisco, México
Izeni Pires Farias Laboratório de Evolução e Genética Animal, Departamento de
Genética, Instituto de Ciências Biológicas, Universidade Federal do Amazonas,
Jaime Gongora Sydney School of Veterinary Science, Faculty of Science,
University of Sydney, Sydney, NSW, Australia
Sandra Marcela Hernández-Rangel Departamento de Genética, Universidade
Federal do Amazonas, Manaus, Amazonas, Brasil
Tomas Hrbek Departamento de Genética, Universidade Federal do Amazonas,
Sally R. Isberg Centre for Crocodile Research, Noonamah, Northern Territory,
Australia
xi
xii Contributors
Igor Joventino Roberto Departamento de Genética, Universidade Federal do

Amazonas, Manaus, Amazonas, Brasil
William E. Magnusson Instituto Nacional de Pesquisas da Amazônia, Manaus,
AM, Brasil
Gualberto Pacheco-Sierra Laboratorio de Genética, Departamento de Ciencias
Naturales, Facultad de Humanidades y Ciencias, Universidad Nacional del Litoral,
Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Santa Fe,
Argentina
Carlos I. Piña CICYTTP (CONICET-Prov. ER-UADER), Facultad de Ciencia y
Tecnología, Diamante, Entre Ríos, Argentina
David A. Ray Department of Biological Sciences, Texas Tech University,
Lubbock, TX, USA
Karina L. Silva-Brandão Centro de Ciências Naturais e Humanas, Universidade
Federal do ABC, Santo André, SP, Brazil
Melina Simoncini CICYTTP (CONICET-Prov. ER-UADER), Facultad de Ciencia
y Tecnología, Diamante, Entre Ríos, Argentina
Pablo Siroski Laboratorio de Ecología Molecular Aplicada-ICiVet UNL/
CONICET, Santa Fe, Argentina
Luciano M. Verdade Universidade de São Paulo, CENA/LE²Ave, Piracicaba, São
Paulo, Brasil
Francisco Villamarín Universidad Regional Amazónica Ikiam, Tena, Ecuador
Rodrigo Barban Zucoloto Laboratório de Genética de Populações e Evolução
Molecular (GENEV), Instituto de Biologia, Universidade Federal da Bahia,
Salvador, Bahia, Brasil
About the Editors
Rodrigo Barban Zucoloto Universidade Federal da Bahia, Brasil

Dr. Zucoloto has a Graduate degree in Biological Sciences from the University
of Sao Paulo (1994), Master of Sciences (Nuclear Energy in Agriculture) from the
University of Sao Paulo (1999), and Doctor of Sciences (Nuclear Energy in
Agriculture) from the University of Sao Paulo (2003). Currently he is Associate
Professor at Federal University of Bahia. Dr. Zucoloto has experience in genetics,
with an emphasis in conservation genetics of Brazilian crocodilians, especially the
broad-snouted caiman (Caiman latirostris).
Patricia Susana Amavet Universidad Nacional del Litoral, Argentina

Dr. Amavet has a Ph.D. in Biological Sciences (2009) from the University of
Buenos Aires. Currently, she is a Professor of Genetics and Population Genetics at
the University National of Litoral. She is a CONICET Researcher since 2010,
Coordinator of the Genetics Laboratory at FHUC-UNL, and Supervisor of three
doctoral students and of one postdoc. She has extensive experience in conservation
genetics of wild species, especially the broad-snouted caiman (Caiman latirostris),
and she is a member of the Crocodile Specialists Group (SSC/IUCN) since 2002.
Her research works include the topics of molecular ecology, population genetics,
mating system studies, and phylogeographic analyses, mainly in reptiles and
amphibians.
Luciano Martins Verdade Universidade de São Paulo, Brasil

Dr. Verdade is an Agronomist and received his Master’s degree in Agronomy
from the University of Sao Paulo (1985 and 1992, respectively) and Ph.D. in
Wildlife Ecology and Conservation from the University of Florida (1997). He is
currently Associate Professor III at the Center for Nuclear Energy in Agriculture
(CENA), University of São Paulo (USP), and member of the coordination of the
Biota / FAPESP Program, the Sao Paulo Biodiversity Commission and the Species
Survival Commission / IUCN, and Information and Management System for
Protected Areas (SIGAP) / Secretariat of Environment of the State of Sao Paulo
(SMA / SP). His research is focused on Applied Evolutionary Ecology with special
xiii
xiv About the Editors
interest in fauna management in agricultural landscapes and adaptive processes of

vertebrates to anthropic alterations.
Izeni Pires Farias Universidade Federal do Amazonas, Brasil

Dr. Farias holds a Bachelor’s degree in Biological Sciences from the Federal
University of Amazonas (1987), Master’s degree in Freshwater Biology and Inner
Fishing from the National Institute of Amazonian Research (1992), and Ph.D. in
Biological Sciences (Molecular Genetics) from Federal University of Pará (2000).
She was a Postdoctoral Fellow (CNPq 2015) at Brigham Young University and is
currently Full Professor at the Federal University of Amazonas where she coordi-
nates the Laboratory of Evolution and Animal Genetics (LEGAL). She has experi-
ence in genetics, with emphasis on population genetics, conservation genetics,
molecular ecology, and phylogeography of Amazonian organisms.
Chapter 1
Geographic Distribution, Habitat,
Reproduction, and Conservation Status
of Crocodilians in the Americas
Francisco Villamarín, Armando H. Escobedo-Galván, Pablo Siroski,

and William E. Magnusson
Abstract The chapter gives an introductory overview on the biology of the 11 spe-
cies of crocodilians inhabiting the Americas. Geographical distribution is broadly
discussed in the light of biogeographical and evolutionary origins. As a broad gen-
eralization, crocodilians are morphologically, genetically, and ecologically conser-
vative and show a high dispersal ability, which together complicate the delimitation
of species boundaries. Furthermore, distinct clades of crocodilians are a result of
environmental selection for different adaptive characteristics. Gene flow occurs
between clades, but distinct morphological varieties are sometimes a result of envi-
ronmental filters. Thus, many taxa of crocodilians can be considered ecological
species. Reproductive characteristics, such as nest-site choice and female reproduc-
tive output, influence reproductive success and thus population trends, so we dis-
cuss reproductive and habitat characteristics in the light of conservation genetics.
Finally, the status of crocodilians is discussed both in terms of past hunting pressure
and current conservation and management initiatives. In general, crocodilians can
look after themselves if they have sufficient habitat, but some species are critically
endangered by hunting, even though much of their original habitat remains intact.
Conservation actions must be applied locally, taking into account threats and
dispersal between source and sink populations. Given the difficulty of studying the
F. Villamarín (*)
Grupo de Biogeografía y Ecología Espacial (BioGeoE2). Universidad Regional Amazónica
Ikiam, Km 7 vía Muyuna., Tena, Ecuador
A. H. Escobedo-Galván
Centro Universitario de la Costa, Universidad de Guadalajara,
Puerto Vallarta, Jalisco, Mexico
P. Siroski
Laboratorio de Ecología Molecular Aplicada-ICiVet UNL/ CONICET, Santa Fe, Argentina
e-mail: [email protected]
W. E. Magnusson
Instituto Nacional de Pesquisas da Amazônia, Manaus, AM, Brazil
e-mail: [email protected]
© Springer Nature Switzerland AG 2021 1

R. B. Zucoloto et al. (eds.), Conservation Genetics of New World Crocodilians,
https://doi.org/10.1007/978-3-030-56383-7_1
2 F. Villamarín et al.
dispersal of crocodilians by direct observation in most places in Latin America, the

definition of effective management units will depend on genetic studies that can be
linked to economic or conservation goals.
Keywords Species boundaries · Ecological species · Reproductive output ·

Mating systems · Conservation strategies
1.1 Introduction
Crocodilians belong to an ancient group that has diversified morphologically and

genetically very little in comparison to other archosaurs, such as dinosaurs, even if
you only consider the only extant lineage, the birds. Other groups that originated
about the same time, such as lizards and mammals, also show much greater mor-
phological diversity. Although some crocodilian lineages produced terrestrial forms,
all extant crocodilians are semiaquatic predators that hunt around the shorelines of
rivers, lakes, estuaries, swamps, and marshes (Grigg and Kirshner 2015).
Demographically, crocodilians are also different from most other vertebrates,
which can generally be categorized as R (short life span, many offspring, usually
small size) or K (long life, often large size, few offspring) selected. Crocodilians are
long lived and large sized, invest heavily in parental care, but produce many off-
spring that are small in relation to the parents (Magnusson 1986). Large males often
sire a disproportionate number of offspring, so effective population sizes may be
much smaller than the number of individuals (Grigg and Kirshner 2015).
The large number of offspring, which usually disperse during the 10 years or
more of adolescence, means that crocodilians have high intrinsic rates of increase
and they tend to colonize all suitable habitat relatively quickly. However, they gen-
erally appear to be limited by low temperatures, and few species (e.g., Alligator
mississippiensis, Alligator sinensis, Crocodylus niloticus) extend beyond the tropics
and subtropics. Alligatorids also lack effective salt secretion glands, and they spend
much less time in saline environments than similar-sized crocodylids, possibly lim-
iting long-distance marine dispersal (Grigg and Kirshner 2015).
Being morphologically, genetically, and ecologically conservative, combined
with high dispersal ability, complicates the definition of species boundaries in croc-
odilians. Vicariance events may cause differentiation of lineages, but many of these,
at least within genera, retain the capacity to produce fertile offspring if brought
together in captivity. Therefore, gene flow between clades that have been isolated
for millions of years may occur if geographic barriers are breached naturally or by
introductions (Nguyen et al. 2018).
It is easy to detect geographically restricted clades of crocodilians that are genet-
ically distinct, but as these clades can usually interbreed if brought into contact, the
question arises as to whether those differences just reflect genetic drift or local
adaptations. This quandary does not only apply to crocodilians. For instance, within
1 Geographic Distribution, Habitat, Reproduction, and Conservation Status… 3
recent historical times, most of the population of humans on the Indian subcontinent
shared mitochondrial DNA almost completely restricted to that region (Reich 2018).
However, nobody is suggesting that the people there represented a distinct species.
Species boundaries are often difficult to define in plants, and many species con-
fined to different habitats produce fertile hybrids at habitat boundaries (Ladiges
1997). These can be considered ecological species, the distinct clades being a result
of environmental selection for different adaptive characteristics. Something similar
seems to occur in many species pairs of crocodylids, one species predominating in
saline environments and the other in adjacent freshwater habitats. Gene flow occurs
between them, but environmental filters apparently result in distinct morphological
varieties at the extremes of salinity (Pacheco-Sierra et al. 2018). The best conserva-
tion strategy for maintaining this genetic variability may not be immediately
obvious.
In this chapter, we discuss aspects on geographic distributions, habitat, reproduc-
tion, and conservation status relevant for the conservation genetics of New World
crocodilians.
1.2 Geographic Distribution
Geographic delimitation of variation in crocodilians is essential to accurately pre-

serve the natural diversity of independent evolutionary lineages. Besides the funda-
mental nature of species as separately evolving genetic lineages, species generally
have unique ecological, morphological, phylogenetic, and life history characteris-
tics that can be used not only for taxonomic identity but to define geographic distri-
bution (De Queiroz 2007; Leaché et al. 2009; Barrett and Freudenstein 2011). To
date, the factors proposed to explain the geographic distribution of the order
Crocodylia around tropical and subtropical regions in the Americas were oversim-
plified mainly due to limitations of theoretical and fieldwork techniques
(Groombridge 1987; Thorbjarnarson et al. 1992; Ross 1998, see Fig. 1.1).
Fossil crocodilian records are fragmented and insufficient to understand biogeo-
graphic processes and ecological relationships. The extant diversity of crocodilians
is not homogeneously represented in fossil records. The genus Crocodylus in the
Americas is not fully represented in the fossil record, and, for instance, we are not
aware of any study reporting fossils of Crocodylus intermedius. Only one record has
been assigned to Crocodylus moreletii of the Pleistocene from Guatemala (Mook
et al. 1959), while other fossil records from Miocene-Pleistocene remain as
Crocodylus sp. (Mead et al. 2006b). Fossil records show that the geographic distri-
butions of Crocodylus acutus and C. rhombifer were broader in the Pleistocene. The
latter belongs mainly to the Caribbean islands (Varona 1984; Morgan and Albury
2013) and C. acutus around the Gulf of California (Mead and Baez 2003; García
2004; Mead et al. 2006a; White et al. 2010; Cupul-Magaña et al. 2017).
The evolutionary origins of alligatorids trace back to the Cretaceous (Brochu
1999; Oaks 2011), and there are good fossil records of the American alligator
Fig. 1.1 Geographic distribution of crocodilians in the Americas based on IUCN Red List infor-
mation available
(Alligator mississippiensis) (Brochu 1999). Some fossils of the genus Caiman

belong to the Miocene and/or Pleistocene from Argentina (Barrios 2013; Bona et al.
2014), Brazil (Fortier et al. 2014), Venezuela (Fortier and Rincón 2013), and Mexico
(Brochu and Carbot-Chanona 2015). Recently, Cidade et al. (2019) examined and
concluded that Caiman venezuelensis (Fortier and Rincón 2013) is a junior syn-
onym of Caiman crocodilus; therefore, it is the first record assigned to this species.
Few records involve material of the genera Melanosuchus and Paleosuchus (Bona
et al. 2017), which are insufficient to determine whether they are closely related or
if the distribution of both genera has been restricted to South America since the late
Miocene.
Despite the limitations of the fossil record, new analytical tools, such as geo-
metrical morphology, molecular techniques, and analytical methods have provided
challenges to the classical view of the definition of crocodilian species boundaries,
with implications for systematics and conservation within the order Crocodylia.
Some studies show that there are likely cryptic lineages, which further complicate
the interpretation of their geographic distribution (McAliley et al. 2006; Venegas-
Anaya et al. 2008; Eaton et al. 2009; Hekkala et al. 2011; Milián-García et al. 2011;
Shirley et al. 2014; Pacheco-Sierra et al. 2018).
Understanding crocodilian distributions in biogeographic studies provides a
basis to identify essential requirements for biological performance of species
(Kadmon et al. 2003; Araujo and Guisan 2006). Determining which factors con-
strain species’ distributions is difficult due to distinct geographical variations and
scales of analysis. In addition, geographic distributions of crocodilians overlap;
thus, geographic distributions, not to mention species delimitation, are difficult to

predict and/or quantify. Overlapping areas imply some ecological and evolutionary
processes. Available information shows that contact areas between species of differ-
ent genera are common. For instance, Alligator and Crocodylus in Florida (Mazzotti
1983), Crocodylus and Caiman from Chiapas, Mexico, to South America (Bolaños
et al. 1996; Martínez-Ibarra et al. 1997; Espinosa-Blanco and Seijas 2010), and
among Melanosuchus, Caiman, and Paleosuchus in the Amazon basin (Medem
1983; Ouboter 1996; Marioni et al. 2008). This results in a similitude of spatial
areas (sympatric species), but niche segregation of syntopic species, which leads to
different spatial and temporal patterns of habitat use (e.g., Ouboter 1996), or diet
partitioning (Villamarín et al. 2017) in mixed populations.
Overlapping areas within the genera Crocodylus, Caiman, and Paleosuchus have
resulted in areas of hybridization and genetic introgression between sympatric spe-
cies. Ray et al. (2004) show the presence of haplotypes characteristic of the
American crocodile (Crocodylus acutus) among some populations of Morelet’s
crocodile (C. moreletii) in Belize, indicating hybridization between the two species
in natural conditions (Fig. 1.2). These results triggered subsequent genetic studies
not only on these species (Cedeño-Vázquez et al. 2008; Rodriguez et al. 2008;
Machkour-M’Rabet et al. 2009; González-Trujillo et al. 2012) but on the potential
hybridization between the C. acutus and the Cuban crocodile (Crocodylus rhombi-
fer) and between C. acutus and the Orinoco crocodile (C. intermedius) (Venegas-
Anaya et al. 2008; Weaver et al. 2008; Milián-García et al. 2011). Recently, genomic
analyses within and between C. acutus and C. moreletii demonstrate that this
hybridization system challenges the definition of species boundaries: C. acutus
could be two distinct species, while C. moreletii might eventually be driven to local
population extirpations (Pacheco-Sierra et al. 2018). The overlapping areas between
C. acutus and C. moreletii showed that the hybridization and/or introgression areas
are wider than previously thought. In addition, the impact of transformed habitat
and hybridization areas among Crocodylus species decreases their potential distri-
bution areas (e.g., Escobedo-Galván and González-Salazar 2011).
In the Amazon basin, Hrbek et al. (2008) evaluated the phylogenetic relation-
ships of the Amazonian caiman species in the Madeira River drainage. The authors
observed that the spectacled caiman (Caiman crocodilus) and the yacare caiman
(C. yacare) share mitochondrial and nuclear haplotypes, while the Schneider’s
dwarf caiman (Paleosuchus trigonatus) and the Cuvier’s dwarf caiman (P. palpe-
brosus) do not, in spite of their phenotypic similarity. Recent molecular analysis
also revealed that the black caiman (Melanosuchus niger) is far more diverse than
previously thought, suggesting the potential existence of different lineages (De
Thoisy et al. 2006; Vasconcelos et al. 2008). Furthermore, Borges et al. (2018) pro-
posed the existence of at least three evolutionary significant units for broad-snouted
caiman (Caiman latirostris).
The use of species distribution models, based on ecological niche model con-
cepts, to predict potential distribution areas has shown that the potential geographic
distribution of some species is broader than previously thought. For instance,
Escobedo-Galván and González-Salazar (2011) showed that the potential
Fig. 1.2 (a) Crocodylus acutus. (b) Crocodylus moreletii. (Photo: Marco A. López-Luna)
distribution of C. acutus includes some areas of the Yucatan Peninsula and the state
of Tabasco, where this species has been considered absent. For C. moreletii, the
distribution models included areas from the southern region of the state of
Tamaulipas up to northern Honduras. If confirmed, this would extend the current
distribution range of C. moreletii at least 50 km to the south. Caiman crocodilus is
widely distributed from Southern Mexico, through Mesoamerica, to northern South
America, and to the Amazon basin (Groombridge 1987; Velasco and Ayarzagüena
2010). This is the most diverse crocodilian species, exhibiting a large morphological
variation through its distributional range, which possibly reflects ecological adapta-
tions to local conditions, suggesting the existence of evolutionary independent lin-
eages recognized currently as C. c. fuscus, C. c. chiapasius, C. c. apaporiensis, and
C. c. crocodilus (Venegas-Anaya et al. 2008; Velasco and Ayarzagüena 2010), or
possibly new lineages previously overlooked (Angulo-Bedoya et al. 2019).
Compared with other crocodilian species, its distribution is well known, but little
attention has been focused on exploring whether biotic and/or abiotic factors might
constrain its distribution. One of the most common questions is related with its
northernmost latitude limit. The northernmost locality of C. crocodilus is consistent
with the type locality of C. crocodilus chiapasius described by Bocourt (1876) from
Isthmus of Tehuantepec, Mexico. The Tehuantepec region is a geographical barrier
to a variety of faunal groups, such as birds, mammals, and butterflies (Peterson et al.
1999; García-Moreno et al. 2004; Barber and Klicka 2010). However, few studies
have focused on evaluating the abiotic parameters that are involved in this geo-
graphical barrier and hence how the Isthmus of Tehuantepec may play a role as an
environmental barrier to the dispersal of some species. Thus, Serna-Lagunes et al.
(2017) conducted species distribution modeling to identify possible climate param-
eters limiting C. crocodilus distributions. Their results show that there are optimal
environmental conditions in the states of the Mexican Pacific coast and in some
areas of the Caribbean. However, the distribution in Mexico of C. crocodilus is
restricted to the coast of Chiapas (Escobedo-Galván et al. 2015). Therefore, its geo-
graphic distribution could be constrained by other factors.
Some species distribution models have been conducted to evaluate the potential
areas of occupation at regional scales in some countries or ecoregions. For example,
the distribution of Caiman yacare in Bolivia (Rodriguez-Cordero et al. 2017),
Crocodylus rhombifer in Cuba (Ramos-Targarona et al. 2008), C. acutus in Ecuador
(Reyes-Puig et al. 2017), C. moreletii in Mexico (Sánchez Herrera et al. 2011), and
C. intermedius in Colombia and Venezuela (Balaguera-Reina et al. 2017). Even if
these efforts allow inferences about local distributions and establishing their conser-
vation and population status, many missing pieces need to be in place before it is
possible to understand the geographic distribution of crocodilians of the Americas.
Future work should address questions regarding how the distribution limits of croc-
odilians could be constrained by environmental conditions or how interspecific
interactions between crocodilians could explain distribution limits when environ-
mental factors are unclear.
1.3 Habitat
As a broad generalization, extant crocodilians are semiaquatic reptiles that thrive in

rivers, small streams, lakes, estuaries, swamps, and marshes. Habitat use of sympat-
ric crocodilians may be very complex and often involves the dominance of larger
and more aggressive species for the most suitable habitats over smaller or less
aggressive species. However, preferential access to suitable sites may not always
involve agonistic encounters. Possibly, as a result of competition, each species gen-
erally occupies different areas and uses available habitats in different ways
(Lang 1987).
Spatial segregation is a common feature among crocodilians. For example,
although Caiman yacare and C. latirostris (Fig. 1.3) are both generalists and occupy
a wide spectrum of habitat types, in places where the species occur sympatrically,
the latter is generally found in still waters with more dense vegetation (Medem
1983) and more ephemeral habitats (Scott Jr. et al. 1990) than those occupied by
C. yacare. In the Río Cojedes in Venezuela, Caiman crocodilus and Crocodylus
intermedius coexist in the same waterway, but generally use different microhabitats.
The latter is mainly found in open waters and within emergent aquatic vegetation
distant from the shore, whereas C. crocodilus mainly occupies water bodies outside
the main channel of the river (Espinosa-Blanco and Seijas 2010).
Thus, despite overlapping geographic distributions and an overall generalist
behavior, some life history traits, such as habitat use, prey preferences, or body size,
may minimize competition between sympatric species (Grigg and Kirshner, 2015).
The geographic distributions of the four species of Amazonian caimans overlap
to a large degree, and they are sometimes found in the same water bodies (Marioni
et al. 2013). However, these species show habitat preferences and are most com-
monly found occupying specific water-body types. Melanosuchus niger occupies
large rivers, streams, lakes, and seasonally flooded savannas, but floodplain lakes
appear to be its main habitat type (Thorbjarnarson 2010). At the other extreme,
Paleosuchus trigonatus is the only caiman typically found occupying small forest
streams that drain large tracts of rainforest (Magnusson 1992; Fig. 1.4). Paleosuchus
palpebrosus most commonly inhabits flooded forests near major rivers and lakes
(Magnusson 1985), Mauritia palm swamps (Godshalk 1982), and streams lined by
gallery forests (Thorbjarnarson et al. 1992; Fig. 1.5). Caiman crocodilus is the most
versatile caiman species as to habitat use. It occurs in high densities in different
habitat types, including preserved wetlands to man-made habitats (Velasco and
Ayarzagüena 2010). Differences in diet among these species may be influenced by
the availability of different prey among habitat types. Since many species of croco-
dilians exhibit ontogenetic shifts in diet (Dodson 1975; Magnusson et al. 1987;
Erickson et al. 2003), individuals of different sizes may occupy different habitats
according to the availability of specific prey (Rosenblatt and Heithaus 2011).
In general, crocodilians have been widely regarded as opportunistic and general-
ist predators. However, studies on A. mississippiensis have shown diet specializa-
tions at the individual level (Rosenblatt et al. 2015), which have cast some doubt on
Fig. 1.3 (a) Caiman latirostris. (b) Caiman yacare. (Photo: Pablo Siroski)
Fig. 1.4 Paleosuchus trigonatus in its typical habitat, next to a small forest stream. (Photo:
Francisco Villamarín)
Fig. 1.5 Paleosuchus palpebrosus inside a flooded forest stream. (Photo: Boris Marioni)
the designation of all crocodilians as generalist predators in all situations. For exam-
ple, the two species of dwarf caimans (P. palpebrosus and P. trigonatus) might avoid
competition by partitioning their diet in places where they occur syntopically
(Villamarín et al. 2017). This suggests that crocodilian divergences in diet might
result from prey preferences and not only from habitat selection.
Differences in habitat selection and foraging behavior could reduce the potential
for competition. For example, A. mississippiensis inhabits swamps, lakes, marshes,
streams, and rivers and may even be found in brackish water (Elsey and Woodward
2010). Crocodylus acutus usually occurs in marine or brackish-water environments,
though it also occasionally occupies freshwater inland areas (Thorbjarnarson 2010).
In situations where two species of the genus Crocodylus come into contact, hybrid-
ization may occur in transitional habitats. For example, hybridization of C. acutus
with the freshwater species C. rhombifer is common in Cuba (Ramos 2008; Ramos-
Targarona et al. 2010). In Belize and Mexico, hybridization has been reported
between C. acutus and C. moreletii (Hekkala 2004; Ray et al. 2004; Cedeño-
Vázquez et al. 2008; Rodriguez et al. 2008; Hekkala et al. 2015; Pacheco-Sierra
et al. 2016). The latter species inhabits mainly freshwater areas such as marshes,
swamps, ponds, rivers, lagoons, and man-made water bodies, but occasionally is
found in brackish or saline habitats, as well (Álvarez del Toro 1974; Escobedo-
Galvan et al. 2008; Platt et al. 2008).
There is evidence that habitat selects for different crocodilian species rather than
the species selecting different habitats. Within the distribution of the genus
Crocodylus, physiological specialization possibly restricts the principal morphot-
ypes to either fresh or saline water in the Caribbean and perhaps in other areas. The
recognized taxa are essentially ecological species, the environment maintaining the
morphotypes distinct at the limits of the environmental cline despite continuous
genetic flux. This is more similar to the situation found in plants than in most verte-
brate genera. In fact, some lineages within the taxa recognized as Crocodylus acutus
are more closely related to other taxa that are generally found in freshwater, such as
C. moreletii or C. intermedius, than they are to each other (Milián-García et al.
2018). It is not clear why the morphology reflects more the environment than phy-
logeny, but it could be related to linked genes or strong selection for a given mor-
photype in each habitat (e.g., Labarre et al. 2017). Freshwater and saline
environments tend to differ in the density of aquatic macrophytes, and it is possible
that the generally broader snouts in freshwater species result from selection for
capturing prey in dense vegetation (Magnusson 2017).
Spatial segregation and different diet preferences when within the same water
body are therefore some of many ways that could allow crocodilians to avoid com-
petitive exclusion and facilitate their coexistence.
1.4 Reproduction
All members of the order Crocodylia share old and conservative reproductive fea-
tures, such as internal fertilization, egg laying, and an elaborate reproductive behav-
ior, including parental care (reviewed by Grigg and Kirshner 2015). These features
are shared with the only other extant archosaur group, the birds.
Reproduction involves complex processes from mating, nesting, and egg incuba-
tion to maternal care of eggs and hatchlings. Most of what is known about courtship
and mating behavior of crocodilians from the Americas comes from observations on
captive populations of A. mississippiensis, Caiman crocodilus, Crocodylus more-
letii, C. rhombifer, C. acutus, and C. intermedius (Hunt 1975; Garrick and Lang
1977; Staton and Dixon 1977; Vliet 1987; Thorbjarnarson and Hernández 1993a, b;
Vliet 2001) and field observations on A. mississippiensis (Dinets 2010), Crocodylus
acutus (Alonso Tabet 2009; Charruau and Hénaut 2012), C. moreletii (López-Luna
et al. 2019), and C. rhombifer (Ramos Targarona 2013). Across all crocodilian spe-
cies in which courtship and mating behavior have been studied, the overall patterns
follow those described by Garrick and Lang (1977), which include complex interac-
tions mediated by sensory cues and dominance behavior (reviewed by Senter 2008).
Until relatively recently, the mating systems of all crocodilian species were
thought to be polygynous, where a single large and dominant male inseminates
most of the reproductive females within its territory (Lang 1987). However, multi-
ple paternity in clutches of eggs, first demonstrated for A. mississippiensis (Davis
et al. 2001), is now thought to be a common feature, and the mating systems are
considered to be polygamous across the order Crocodylia. Although no evidence
has been found that suggests higher hatching success in multiple-fathered clutches,
multiple paternity may increase effective population size and genetic diversity and
has been found in many species of crocodilians of the Americas, including Caiman
latirostris (Amavet et al. 2008, 2012), C. crocodilus (Oliveira et al. 2014), C. yacare
(Ojeda et al. 2016), Melanosuchus niger (Muniz et al. 2011), Crocodylus interme-
dius (Lafferriere et al. 2016), C. moreletii (McVay et al. 2008), and C. rhombifer
(Milián-García et al. 2016).
Most studies on the reproduction of Neotropical crocodilians have focused on
nesting and the reproductive output of females. Together, these are important life
history features that may influence demographic trends. In a comprehensive analy-
sis of the reproductive characteristics of the order Crocodylia, Thorbjarnarson
(1996) found that, overall, species in the family Alligatoridae produce larger
clutches but reproduce less frequently than those in the Crocodylidae. These differ-
ent reproductive strategies lead to similar levels of reproductive output in the two
families. Within the Alligatoridae, there are a few differences in the reproductive
characteristics among genera. Alligator mississippiensis, the only alligatorid in the
Americas living in temperate regions, produces large clutches of small eggs but with
a low annual clutch frequency. On the other hand, clutch mass of alligatorids occur-
ring in tropical regions is much higher as a consequence of large eggs (Paleosuchus)
or large clutches (Caiman) (Thorbjarnarson 1996).
Crocodilians may dig a hole in the ground to lay their eggs or build mounds of
vegetation. Mound nesting is the main mode used by both the Alligatoridae and the
Crocodylidae and presumably is advantageous over hole nesting in most habitats
(Fig. 1.6). The few species that build hole nests usually inhabit seasonal habitats of
rivers or lakes (Campbell 1972), and currently the only crocodilians of the Americas
considered hole nesters are Crocodylus acutus and C. intermedius (reviewed by
Grigg and Kirshner 2015). Some authors have suggested that underground nests
may experience limited gas diffusion (Seymour and Ackerman 1980) and nesting
sites of hole nesting crocodilians might be confined to particular soil characteristics
(Lutz and Dunbar-Cooper 1984; Mazzotti 1989; Casas-Andreu 2003; Gómez-
González et al. 2017).
Another important factor that influences reproductive success in crocodilians is
nest-site selection. Crocodilians presumably nest in sites that present the most
appropriate conditions in relation to factors that influence egg development (e.g.,
temperature, humidity) or egg survival (e.g., flooding, predation) (Webb et al. 1983;
Ouboter and Nanhoe 1988; Somaweera et al. 2011). Crocodilians may exhibit nest-
site fidelity, reusing the same nesting site from previous years. This behavior has
been documented in Alligator mississippiensis (Elsey et al. 2008), Crocodylus acu-
tus (Charruau et al. 2010), and C. moreletii (López-Luna et al. 2011). This might
Fig. 1.6 Mound nest of

Melanosuchus niger in
várzea floodplains in the
Brazilian Amazon. (Photo:
Francisco Villamarín)
reduce the costs associated with selecting suitable nesting sites in subsequent nest-
ing events. Surprisingly, little information is available on behavior plasticity regard-
ing nest-site selection in crocodilians and its benefits for egg viability, hatchling
phenotypes, and individual fitness in wild crocodilian populations (López-Luna
et al. 2020).
Sex determination in crocodilians is temperature-dependent with a female-male-
female pattern (reviewed in Deeming 2004). First discovered in Alligator mississip-
piensis, in general crocodilians produce mostly females at low and high incubation
temperatures, while a majority of males are produced at intermediate temperatures
(Ferguson and Joanen 1982; Deeming 2004; González et al. 2019). Therefore, nest
constructions (hole or mound) are crucial to protect eggs from thermal extremes by
maintaining relatively constant internal temperature regimes (Magnusson et al.
1985, 1990; Villamarín-Jurado and Suárez 2007). Thus, nest-site choice is vital to
ensure the appropriate conditions for the incubation of the eggs and is critical to the
reproductive success of crocodilians. The most productive nesting habitat for most
crocodilian species has been suggested to be freshwater wetlands/grasslands
(Brazaitis and Watanabe 2011). However, an obvious exception among crocodilians
of the Americas appears to be the forest dwelling Paleosuchus trigonatus, which
occurs in small streams most commonly located in terra firme (non-flooded) forests
and their nests are built around those water bodies (Magnusson et al. 1985). As
opposed to most crocodilian species in which eggs are heated by insolation, rotting
vegetation, and metabolic heat of embryos, the internal nest temperature may also
originate from metabolism of termites (Magnusson et al. 1985, 1990).
Another important implication of nest-site selection in crocodilians is the vulner-
ability of eggs and hatchlings to predation. A number of different organisms have
been identified as natural predators of early stages of crocodilians, including inver-
tebrates, fish, anurans, reptiles, birds, and mammals (reviewed in Somaweera et al.
2013). In general, nests located in more exposed sites that have easier access for
predators may face higher levels of predation. For example, nests of C. yacare and
C. c. crocodilus constructed on floating mats and surrounded by water are less prone
to egg predation (Ouboter and Nanhoe 1987; Campos 1993). Similarly, islands may
offer suitable nesting sites as fewer predators occur on islands than on the mainland.
For example, nests of C. moreletii located on natural islands in Belize show higher
hatching success than nests built on the shoreline or on artificial islands (Platt et al.
2008). However, the success of predatory events on crocodilian eggs is also influ-
enced by active parental defense of nests.
Most species of crocodilians actively defend their nests (Shine 1988; Brazaitis
and Watanabe 2011; Fig. 1.7). Although nest defense and territoriality are more
common in mound-nesting species than in hole nesters, aggressive protection of
nests has been documented for most crocodilian species (McIlhenny 1934; Cott
1971; Charruau and Hénaut 2012). Nest defense may influence hatching success.
For example, studies on A. mississippiensis and M. niger have shown that predation
rates on eggs are lower when adults are present (Kushlan and Kushlan 1980;
Torralvo et al. 2017). However, a common cause of egg mortality, often as important
as predation, is the flooding of nests.
Fig. 1.7 (a–c) Camera

trap capture of a
Melanosuchus niger
female guarding its nest.
(d) Aggressive behavior
shown by a Melanosuchus
niger female next to its
nest. (Photo: Francisco
Villamarín)
Fig. 1.7 (continued)
Some crocodilian species generally place their nests very close to the water’s
edge, and adult individuals, presumably females, guard the nests from predators in
the water. Nest guarding of M. niger females might decrease predation, but the
proximity to the water’s edge might increase egg mortality due to flooding. In con-
trast, C. crocodilus females often place their nests hundreds of meters inside the
forest, away from permanent water bodies, and guard hidden under leaf litter
(Villamarín et al. 2011; Barão-Nóbrega et al. 2016, 2018; Fig. 1.8). However, in
such situations, C. crocodilus females are prone to predation by jaguars (Panthera
onca) (Da Silveira et al. 2010). Thus, nest-site selection might result in a trade-off
between higher vulnerability to egg predation when nests are located too far from
the shoreline and flooding risk when nests are too close (Allsteadt 1994; Larriera
and Piña 2000; Villamarín et al. 2011).
1.5 Conservation Status
The conservation status of a wild species is an indicator that tells us the likelihood
of its survivorship at present or in the future. Many factors are considered when
evaluating the conservation status of a species, where the most relevant among
many are the remaining number of individuals, the overall increase or decrease in
the population over time, breeding success rates, and known threats. The conserva-
tion status of a species is ranked in a scientifically based system called IUCN Red
List of Threatened Species, which contains the most comprehensive information on
the global conservation status of animal, fungi, and plant species. The system
divides species into nine categories according to their conservation status: Extinct
(EX), Extinct in the Wild (EW), Critically Endangered (CR), Endangered (EN),
Vulnerable (VU), Near Threatened (NT), Least Concern (LC), Data Deficient (DD),
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The confluence of two rings of equal number of electrons, which
rotate round a nucleus of charge outside a ring of electrons
already bound, must be expected to take place more easily than the
confluence of two similar rings rotating round a nucleus of charge
; for the stability of the rings for a displacement
perpendicular to their plane will (see §2) be smaller in the first than in
the latter case. This tendency for stability to decrease for
displacements perpendicular to the plane of the ring will be especially
marked for the outer rings of electrons of a neutral atom. In the latter
case we must expect the confluence of rings to be greatly facilitated,
and in certain cases it may even happen that the number of electrons
in the outer ring may be greater than in the next, and that the outer
ring may show deviations from the assumption of , , , electrons
in the rings, e. g. the configurations and instead of the
configurations and . We shall here not discuss further
the intricate question of the arrangement of the electrons in the outer
ring. In the scheme given below the number of electrons in this ring is
arbitrarily put equal to the normal valency of the corresponding
element; i. e. for electronegative and electropositive elements
respectively the number of hydrogen atoms and twice the number of
oxygen atoms with which one atom of the element combines.
Such an arrangement of the outer electrons is suggested by
considerations of atomic volumes. As is well known, the atomic
volume of the elements is a periodic function of the atomic weights. If
arranged in the usual way according to the periodic system, the
elements inside the same column have approximately the same
atomic volume, while this volume changes considerably from one
column to another, being greatest for columns corresponding to the
smallest valency and smallest for the greatest valency . An
approximate estimate of the radius of the outer ring of a neutral atom
can be obtained by assuming that the total force due to the nucleus
and the inner electrons is equal to that from a nucleus of charge ,
where is the number of electrons in the ring. Putting
in the equation (1) on p. 28, and denoting the value of for by
, we get for , ; for , ; and for
, . Accordingly the arrangement chosen for the
electrons will involve a variation in the dimensions of the outer ring
similar to the variation in the atomic volumes of the corresponding
elements. It must, however, be borne in mind that the experimental
determinations of atomic volumes in most cases are deduced from
consideration of molecules rather than atoms.
From the above we are led to the following possible scheme for
the arrangement of the electrons in light atoms:—
Without any fuller discussion it seems not unlikely that this

constitution of the atoms will correspond to properties of the elements
similar with those observed.
In the first place there will be a marked periodicity with a period of
. Further, the binding of the outer electrons in every horizontal series
of the above scheme will become weaker with increasing number of
electrons per atom, corresponding to the observed increase of the
electropositive character for an increase of atomic weight of the
elements in every single group of the periodic system. A
corresponding agreement holds for the variation of the atomic
volumes.
In the case of atoms of higher atomic weight the simple
assumptions used do not apply. A few indications, however, are
suggested from consideration of the variations in the chemical
properties of the elements. At the end of the 3rd period of elements
we meet with the iron-group. This group takes a particular position in
the system of the elements, since it is the first time that elements of
neighbouring atomic weights show similar chemical properties. This
circumstance indicates that the configurations of the electrons in the
elements of this group differ only in the arrangement of the inner
electrons. The fact that the period in the chemical properties of the
elements after the iron-group is no longer , but , suggests that
elements of higher atomic weight contain a recurrent configuration of
electrons in the innermost rings. The deviation from , , ,
may be due to a gradual interchange of electrons between the rings,
such as is indicated on p. 45. Since a ring of electrons will not be
stable the electrons may be arranged in two parallel rings (see p. 36).
Such a configuration of the inner electrons will act upon the outer
electrons in very nearly the same way as a nucleus of charge
. It might therefore be possible that with increase of
another configuration of the same type will be formed outside the first,
such as is suggested by the presence of a second period of
elements.
On the same lines, the presence of the group of the rare earths
indicates that for still greater values of another gradual alteration
of the innermost rings will take place. Since, however, for elements of
higher atomic weight than those of this group, the laws connecting
the variation of the chemical properties with the atomic weight are
similar to those between the elements of low atomic weight, we may
conclude that the configuration of the innermost electrons will be
again repeated. The theory, however, is not sufficiently complete to
give a definite answer to such problems.
§5. Characteristic Röntgen Radiation.
According to the theory of emission of radiation given in Part I.,

the ordinary line-spectrum of an element is emitted during the
reformation of an atom when one or more of the electrons in the outer
rings are removed. In analogy it may be supposed that the
characteristic Röntgen radiation is sent out during the settling down of
the system if electrons in inner rings are removed by some agency, e.
g. by impact of cathode particles. This view of the origin of the
characteristic Röntgen radiation has been proposed by Sir J. J.
Thomson[35].
Without any special assumption in regard to the constitution of the
radiation, we can from this view determine the minimum velocity of
the cathode rays necessary to produce the characteristic Röntgen
radiation of a special type by calculating the energy necessary to
remove one of the electrons from the different rings. Even if we knew
the numbers of electrons in the rings, a rigorous calculation of this
minimum energy might still be complicated, and the result largely
dependent on the assumptions used; for, as mentioned in Part I., p.
19, the calculation cannot be performed entirely on the basis of the
ordinary mechanics. We can, however, obtain very simply an
approximate comparison with experiments if we consider the
innermost ring and as a first approximation neglect the repulsion from
the electrons in comparison with the attraction of the nucleus. Let us
consider a simple system consisting of a bound electron rotating in a
circular orbit round a positive nucleus of charge From the expressions
(1) on p. 28 we get for the velocity of the electron, putting ,
The total energy to be transferred to the system in order to

remove the electron to an infinite distance from the nucleus is equal
to the kinetic energy of the bound electron. If, therefore, the electron
is removed to a great distance from the nucleus by impact of another
rapidly moving electron, the smallest kinetic energy possessed by the
latter when at a great distance from the nucleus must necessarily be
equal to the kinetic energy of the bound electron before the collision.
The velocity of the free electron therefore must be at least equal to .
According to Whiddington’s experiments[36] the velocity of
cathode rays just able to produce the characteristic Röntgen radiation
of the so-called -type—the hardest type of radiation observed—
from an element of atomic weight is for elements from Al to Se
approximately equal to . As seen this is equal to the
above calculated value for , if we put .
Since we have obtained approximate agreement with experiment

by ascribing the characteristic Röntgen radiation of the -type to the
innermost ring, it is to be expected that no harder type of
characteristic radiation will exist. This is strongly indicated by
observations of the penetrating-power of rays[37].
It is worthy of remark that the theory gives not only nearly the right
value for the energy required to remove an electron from the outer
ring, but also the energy required to remove an electron from the
innermost ring. The approximate agreement between the calculated
and experimental values is all the more striking when it is recalled
that the energies required in the two cases for an element of atomic
weight differ by a ratio of .
In connexion with this it should be emphasized that the
remarkable homogeneity of the characteristic Röntgen radiation—
indicated by experiments on absorption of the rays, as well as by the
interference observed in recent experiments on diffraction of Röntgen
rays in crystals—is in agreement with the main assumption used in
Part I. (see p. 7) in considering the emission of line-spectra, viz. that
the radiation emitted during the passing of the systems between
different stationary states is homogeneous.
Putting in (4) , we get for the diameter of the innermost
ring approximately . For this gives
, a value which is very small in comparison with
ordinary atomic dimensions but still very great compared with the
dimensions to be expected for the nucleus. According to Rutherford’s
calculation the dimensions of the latter are of the same order of
magnitude as .
§6. Radioactive Phenomena.

According to the present theory the cluster of electrons
surrounding the nucleus is formed with emission of energy, and the
configuration is determined by the condition that the energy emitted is
a maximum. The stability involved by these assumptions seems to be
in agreement with the general properties of matter. It is, however, in
striking opposition to the phenomena of radioactivity, and according
to the theory the origin of the latter phenomena may therefore be
sought elsewhere than in the electronic distribution round the
nucleus.
A necessary consequence of Rutherford’s theory of the structure
of atoms is that the -particles have their origin in the nucleus. On
the present theory it seems also necessary that the nucleus is the
seat of the expulsion of the high-speed -particles. In the first place,
the spontaneous expulsion of a -particle from the cluster of
electrons surrounding the nucleus would be something quite foreign
to the assumed properties of the system. Further, the expulsion of an
-particle can hardly be expected to produce a lasting effect on the
stability of the cluster of electrons. The effect of the expulsion will be
of two different kinds. Partly the particle may collide with the bound
electrons during its passing through the atom. This effect will be
analogous to that produced by bombardment of atoms of other
substances by -rays and cannot be expected to give rise to a
subsequent expulsion of -rays. Partly the expulsion of the particle
will involve an alteration in the configuration of the bound electrons,
since the charge remaining on the nucleus is different from the
original. In order to consider the latter effect let us regard a single ring
of electrons rotating round a nucleus of charge , and let us
assume that an -particle is expelled from the nucleus in a direction
perpendicular to the plane of the ring. The expulsion of the particle
will obviously not produce any alteration in the angular momentum of
the electrons; and if the velocity of the -particle is small compared
with the velocity of the electrons—as it will be if we consider inner
rings of an atom of high atomic weight—the ring during the expulsion
will expand continuously, and after the expulsion will take the position
claimed by the theory for a stable ring rotating round a nucleus of
charge . The consideration of this simple case strongly
indicates that the expulsion of an -particle will not have a lasting
effect on the stability of the internal rings of electrons in the residual
atom.
The question of the origin of -particles may also be considered
from another point of view, based on a consideration of the chemical
and physical properties of the radioactive substances. As is well
known, several of these substances have very similar chemical
properties and have hitherto resisted every attempt to separate them
by chemical means. There is also some evidence that the substances
in question show the same line-spectrum[38]. It has been suggested
by several writers that the substances are different only in radioactive
properties and atomic weight but identical in all other physical and
chemical respects. According to the theory, this would mean that the
charge on the nucleus, as well as the configuration of the surrounding
electrons, was identical in some of the elements, the only difference
being the mass and the internal constitution of the nucleus. From the
considerations of §4 this assumption is already strongly suggested by
the fact that the number of radioactive substances is greater than the
number of places at our disposal in the periodic system. If, however,
the assumption is right, the fact that two apparently identical elements
emit -particles of different velocities, shows that the -rays as well
as the -rays have their origin in the nucleus.
This view of the origin of - and -particles explains very simply
the way in which the change in the chemical properties of the
radioactive substances is connected with the nature of the particles
emitted. The results of experiments are expressed in the two
rules[39]:—
1. Whenever an -particle is expelled the group in the periodic
system to which the resultant product belongs is two units less than
that to which the parent body belongs.
2. Whenever a -particle is expelled, the group of the resultant
body is unit greater than that of the parent.
As will be seen this is exactly what is to be expected according to
the considerations of §4.
In escaping from the nucleus, the -rays may be expected to
collide with the bound electrons in the inner rings. This will give rise to
an emission of a characteristic radiation of the same type as the
characteristic Röntgen radiation emitted from elements of lower
atomic weight by impact of cathode-rays. The assumption that the
emission of -rays is due to collisions of -rays with bound electrons
is proposed by Rutherford[40] in order to account for the numerous
groups of homogeneous -rays expelled from certain radioactive
substances.
In the present paper it has been attempted to show that the

application of Planck’s theory of radiation to Rutherford’s atom-model
through the introduction of the hypothesis of the universal constancy
of the angular momentum of the bound electrons, leads to results
which seem to be in agreement with experiments.
In a later paper the theory will be applied to systems containing
more than one nucleus.
[23] Communicated by Prof. E. Rutherford, F.R.S.
[24] Part I. was published in Phil. Mag. xxvi. p. 1 (1913).
[25] Comp, also Geiger and Marsden, Phil. Mag. xxv. p. 604
(1913).
[26] Comp. C. G. Barkla, Phil. Mag. xxi. p. 648 (1911).
[27] Comp. A. v. d. Broek, Phys. Zeitschr. xiv. p. 32 (1913). Phil.
Mag. S. 6. Vol. 26. No. 153. Sept. 1913.
[28] Comp. J. W. Nicholson, Month. Not. Roy. Astr. Soc. 72.
p.52(1912).
[29] This value is that calculated in the first part of the paper.
Using the values (see R. A. Millikan, Brit.
Assoc. Rep. 1912, p. 410), (see P. Gmelin,
Ann. d. Phys. xxviii. p. 1086 (1909) and A. H. Bucherer, Ann. d.
Phys. xxxvii. p. 597 (1912)),and (calculated by
Planck’s theory from the experiments of E. Warburg, G.
Leithäuser, E. Hupka, and C. Müller, Ann. d. Phys. xl. p. 611
(1913)) we get in very close agreement

with observations.
[30] J. J. Thomson, Phil. Mag. xxiv. p. 218 (1912).
[31] J. Franck u. G. Hertz, Verh. d. Deutsch. Phys. Ges. xv. p. 34
(1913).
[32] C. and M. Cuthbertson, Proc. Roy. Soc. A. lxxxiv. p. 13
(1910). (In a previous paper (Phil. Mag. Jan. 1913) the author
took the values for the refractive index in helium, given by M. and
C. Cuthbertson, as corresponding to atmospheric pressure; these
values, however, refer to double atmospheric pressure.
Consequently the value there given for the number of electrons in
a helium atom calculated from Drude’s theory has to be divided by
2.)
[33] See J. Franck, Verh. d. Deutsch. Phys. Ges. xii. p. 613
(1910).
[34] J. W. Nicholson, Month. Not. Roy. Astr. Soc. lxxiii. p. 382
(1913).
[35] Comp. J. J. Thomson, Phil. Mag. xxiii. p. 456 (1912).
[36] R. Whiddington, Proc. Roy. Soc. A. lxxxv. p. 323 (1911).
[37] Comp. E. Rutherford, Phil. Mag. xxiv. p. 453 (1912).
[38] See A. S. Russell and R. Rossi, Proc. Roy. Soc. A. lxxxvii. p.
478 (1912).
[39] See A. S. Russell, Chem. News, cvii. p. 49 (1913); G. V.
Hevesy, Phys. Zeitschr. xiv. p. 49 (1913); K. Fajans, Phys.
Zeitschr. xiv. pp. 131 & 136 (1913): Verh. d. deutsch. Phys. Ges.
xv. p. 240 (1913); F. Soddy, Chem. News, cvii. p. 97 (1913).
[40] E. Rutherford, Phil. Mag. xxiv. pp. 453 & 893 (1912).
Part III.—SYSTEMS CONTAINING
SEVERAL NUCLEI [41] [42].
§1. Preliminary.
ACCORDING to Rutherford's theory of the structure of atoms, the

difference between an atom of an element and a molecule of a
chemical combination is that the first consists of a cluster of electrons
surrounding a single positive nucleus of exceedingly small
dimensions and of a mass great in comparison with that of the
electrons, while the latter contains at least two nuclei at distances
from each other comparable with the distances apart of the electrons
in the surrounding cluster.
The leading idea used in the former papers was that the atoms
were formed through the successive binding by the nucleus of a
number of electrons initially nearly at rest.
Such a conception, however, cannot be utilized in considering the
formation of a system containing more than a single nucleus; for in
the latter case there will be nothing to keep the nuclei together during
the binding of the electrons. In this connexion it may be noticed that
while a single nucleus carrying a large positive charge is able to bind
a small number of electrons, on the contrary, two nuclei highly
charged obviously cannot be kept together by the help of a few
electrons. We must therefore assume that configurations containing
several nuclei are formed by the interaction of systems—each
containing a single nucleus—which already have bound a number of
electrons.
§2 deals with the configuration and stability of a system already
formed. We shall consider only the simple case of a system
consisting of two nuclei and of a ring of electrons rotating round the
line connecting them; the result of the calculation, however, gives
indication of what configurations are to be expected in more
complicated cases. As in the former papers, we shall assume that the
conditions of equilibrium can be deduced by help of the ordinary
mechanics. In determining the absolute dimensions and the stability
of the systems, however, we shall use the main hypothesis of Part I.
According to this, the angular momentum of every electron round the
centre of its orbit is equal to a universal value where is
Planck’s constant; further, the stability is determined by the condition
that the total energy of the system is less than in any neighbouring
configuration satisfying the same condition of the angular momentum
of the electrons.
In §3 the configuration to be expected for a hydrogen molecule is
discussed in some detail.
§4 deals with the mode of formation of the systems. A simple
method of procedure is indicated, by which it is possible to follow,
step by step, the combination of two atoms to form a molecule. The
configuration obtained will be shown to satisfy the conditions used in
§2. The part played in the considerations by the angular momentum
of the electrons strongly supports the validity of the main hypothesis.
§5 contains a few indications of the configurations to be expected
for systems containing a greater number of electrons.
§2. Configurations and Stability of the Systems.
Let us consider a system consisting of two positive nuclei of equal

charges and a ring of electrons rotating round the line connecting
them. Let the number of electrons in the ring be , the charge of an
electron , and the charge on each nucleus . As can be simply
shown, the system will be in equilibrium if the nuclei are the same
distance apart from the plane of the ring and if the ratio between the
diameter of the ring and the distance apart of the nuclei is
given by
provided that the frequency of revolution is of a magnitude such
that for each of the electrons the centrifugal force balances the radial
force due to the attraction of the nuclei and the repulsion of the other
electrons. Denoting this force by , we get from the condition of

the universal constancy of the angular momentum of the electrons, as
shown in Part II. p. 28,
The total energy necessary to remove all the charged particles to

infinite distances from each other is equal to the total kinetic energy
of the electrons and is given by
For the system in question we have
where
a table of is given in Part II. p. 32

To test the stability of the system we have to consider
displacements of the orbits of the electrons relative to the nuclei, and
also displacements of the latter relative to each other.
A calculation based on the ordinary mechanics gives that the
systems are unstable for displacements of the electrons in the plane
of the ring. As for the systems considered in Part II., we shall,
however, assume that the ordinary principles of mechanics cannot be
used in discussing the problem in question, and that the stability of
the systems for the displacements considered is secured through the
introduction of the hypothesis of the universal constancy of the
angular momentum of the electrons. This assumption is included in
the condition of stability stated in §1. It should be noticed that in Part
II. the quantity was taken as a constant, while for the systems
considered here, , for fixed positions of the nuclei, varies with the
radius of the ring. A simple calculation, however, similar to that given
in Part II. on p. 30, shows that the increase in the total energy of the
system for a variation of the radius of the ring from a to ,
neglecting powers of a greater than the second, is given by
where is the total kinetic energy and the potential energy of the
system. Since for fixed positions of the nuclei increases for
increasing , the
term dependent on the variation of will be positive, and the system
will consequently be stable for the displacement in question.
From considerations exactly corresponding to those given in Part
II. on p. 31, we get for the conditional stability for displacements of
the electrons perpendicular to the plane of the ring
where has the same signification as in Part II., and
denotes the component, perpendicular to the plane of the

ring, of the force due to the nuclei, which acts upon one of the
electrons in the ring when it has suffered a small displacement
perpendicular to the plane of the ring. As for the systems considered
in Part II., the displacements can be imagined to be produced by the
effect of extraneous forces acting upon the electrons in direction
parallel to the axis of the system.
For a system of two nuclei each of charge and with a ring of
electrons, we find
By help of this expression and using the table for given

on p. 32 in Part II., it can be simply shown that the system in question
will not be stable unless and equal to or .
In considering the stability of the systems for a displacement of
the nuclei relative to each other, we shall assume that the motions of
the nuclei are so slow that the state of motion of the electrons at any
moment will not differ sensibly from that calculated on the assumption
that the nuclei are at rest. This assumption is permissible on account
of the great mass of the nuclei compared with that of the electrons,
which involves that the vibrations resulting from a displacement of the
nuclei are very slow compared with those due to a displacement of
the electrons. For a system consisting of a ring of electrons and two
nuclei of equal charge, we shall thus assume that the electrons at any
moment daring the displacement of the nuclei move in circular orbits
in the plane of symmetry of the latter.
Let us now imagine that, by help of extraneous forces acting on
the nuclei, we slowly vary the distance between them. During the
displacement the radius of the ring of electrons will vary in
consequence of the alteration of the radial force due to the attraction
of the nuclei. During this variation the angular momentum of each of
the electrons round the line connecting the nuclei will remain
constant. If the distance apart of the nuclei increases, the radius of
the ring will obviously also increase; the radius, however, will increase
at a slower rate than the distance between the nuclei. For example,
imagine a displacement in which the distance as well as the radius
are both increased to a times their original value. In the new
configuration the radial force acting on an electron from the nuclei
and the other electrons is times that in the original configuration.
From the constancy of the angular momentum of the electrons during
the displacement, it further follows that the velocity of the electrons in
the new configuration is times, and the centrifugal force times
that in the original. Consequently, the radial force is greater than the
centrifugal force.
On account of the distance between the nuclei increasing faster
than the radius of the ring, the attraction on one of the nuclei due to
the ring will be greater than the repulsion from the other nucleus. The
work done during the displacement by the extraneous forces acting
on the nuclei will therefore be positive, and the system will be stable
for the displacement. Obviously the same result will hold in the case
of the distance between the nuclei diminishing. It may be noticed that
in the above considerations we have not made use of any new
assumption on the dynamics of the electrons, but have only used the
principle of the invariance of the angular momentum, which is
common both for the ordinary mechanics and for the main hypothesis
of §1.
For a system consisting of a ring of electrons and two nuclei of
unequal charge, the investigation of the stability is more complicated.
As before, we find that the systems are always stable for
displacements of the electrons in the plane of the ring; also an
expression corresponding to (4) will hold for the condition of stability
for displacements perpendicular to the plane of the ring. This
condition, however, will not be sufficient to secure the stability of the
system. For a displacement of the electrons perpendicular to the
plane of the ring, the variation of the radial force due to the nuclei will
be of the same order of magnitude as the displacement; therefore, in
the new configuration the radial force will not be in equilibrium with
the centrifugal force, and, if the radius of the orbits is varied until the
radial equilibrium is restored, the energy of the system will decrease.
This circumstance must be taken into account in applying the
condition of stability of §1. Similar complications arise in the
calculation of stability for displacements of the nuclei. For a variation
of the distance apart of the nuclei not only will the radius of the ring
vary but also the ratio in which the plane of the ring divides the line
connecting the nuclei. As a consequence, the full discussion of the
general case is rather lengthy; an approximate numerical calculation,
however, shows that the systems, as in the former case, will be
unstable unless the charges on the nuclei are small and the ring
contains very few electrons.
The above considerations suggest configurations of systems,
consisting of two positive nuclei and a number of electrons, which are
consistent with the arrangement of the electrons to be expected in
molecules of chemical combinations. If we thus consider a neutral
system containing two nuclei with great charges, it follows that in a
stable configuration the greater part of the electrons must be
arranged around each nucleus approximately as if the other nucleus
were absent; and that only a few of the outer electrons will be
arranged differently rotating in a ring round the line connecting the
nuclei, The latter ring, which keeps the system together, represents
the chemical “bond.”
A first rough approximation of the possible configuration of such a
ring can be obtained by considering simple systems consisting of a
single ring rotating round the line connecting two nuclei of minute
dimensions. A detailed discussion, however, of the configuration of
systems containing a greater number of electrons, taking the effect of
inner rings into account, involves elaborate numerical calculations.
Apart from a few indications given in §5, we shall in this paper confine
ourselves to systems containing very few electrons.
§3. Systems containing few Electrons. The Hydrogen Molecule.

Among the systems considered in §2 and found to be stable the
system formed of a ring of two electrons and of two nuclei of charge
is of special interest, as it, according to the theory, may be expected
to represent a neutral hydrogen molecule.
Denoting the radius of the ring by and the distances apart of the
nuclei from the plane of the ring by , we get from (1), putting
and ,
from (4) we further get
From (2) and (3) we get, denoting as in Part II. the values of , and
for a system consisting of a single electron rotating round a
nucleus of charge (a hydrogen atom) by , , and ,
Since , it follows that two hydrogen atoms combine into a

molecule with emission of energy. Putting
(comp. Part II. p. 38) and , where is the number of
molecules in a gram-molecule, we get for the energy emitted during
the formation of a gram-molecule of hydrogen from hydrogen atoms
, which corresponds to .
This value is of the right order of magnitude; it is, however,
considerably less than the value found by Langmuir[43]
by measuring the heat conduction through the gas from an
incandescent wire in hydrogen. On account of the indirect method
employed it seems difficult to estimate the accuracy to be ascribed to
the latter value. In order to bring the theoretical value in agreement
with Langmuir’s value, the magnitude of the angular momentum of
the electrons should be only ⅔ of that adopted; this seems, however,
difficult to reconcile with the agreement obtained on other points.
From (6) we get . For the frequency of
vibration of the whole ring in the direction parallel to the axis of the
system we get
We have assumed in Part I. and Part II. that the frequency of

radiation absorbed by the system and corresponding to vibrations of
the electrons in the plane of the ring cannot be calculated from the
ordinary mechanics, but is determined by the relation , where
is Planck’s constant, and the difference in energy between two
different stationary states of the system. Since we have seen in §2
that a configuration consisting of two nuclei and a single electron
rotating round the line between them is unstable, we may assume
that the removing of one of the electrons will lead to the breaking up
of the molecule into a single nucleus and a hydrogen atom. If we
consider the latter state as one of the stationary states in question we
get
The value for the frequency of the ultra-violet absorption line in

hydrogen calculated from experiments on dispersion is
.[44] Further, a calculation from such
experiments based on Drude’s theory gives a value near two for the
number of electrons in a hydrogen molecule. The latter result might
have connexion with the fact that the frequencies calculated above
for the radiation absorbed corresponding to vibrations parallel and
perpendicular to the plane of the ring are nearly equal. As mentioned
in Part II., the number of electrons in a helium atom calculated from
experiments on dispersion is only about ⅔ of the number of electrons
to be expected in the atom, viz. two. For a helium atom, as for a
hydrogen molecule, the frequency determined by the relation
agrees closely with the frequency observed from
dispersion; in the helium system, however, the frequency
corresponding to vibrations perpendicular to the plane of the ring is
more than three times as great as the frequency in question, and
consequently of negligible influence on the dispersion.
In order to determine the frequency of vibration of the system
corresponding to displacement of the nuclei relative to each other, let
us consider a configuration in which the radius of the ring is equal to
, and the distance apart of the nuclei . The radial force acting on
one of the electrons and due to the attraction from the nuclei and the
repulsion from the other electron is
Let us now consider a slow displacement of the system during which

the radial force balances the centrifugal force due to the rotation of
the electrons, and the angular momentum of the latter remains
constant. Putting , we have seen on p. 55 that the radius

of the ring is inversely proportional to . Therefore, during the
displacement considered, remains constant. This gives by
differentiation
Introducing and , we get
The force acting on one of the nuclei due to the attraction from the
ring and the repulsion from the other nucleus is

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Keywords Species boundaries · Ecological species · Reproductive output ·

Crocodilians belong to an ancient group that has diversified morphologically and

1.2 Geographic Distribution

Geographic delimitation of variation in crocodilians is essential to accurately pre-

(Alligator mississippiensis) (Brochu 1999). Some fossils of the genus Caiman

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Fig. 1.6 Mound nest of

Fig. 1.7 (a–c) Camera

Fig. 1.7 (continued)

1.5 Conservation Status

Without any fuller discussion it seems not unlikely that this

§5. Characteristic Röntgen Radiation.

According to the theory of emission of radiation given in Part I.,

The total energy to be transferred to the system in order to

Since we have obtained approximate agreement with experiment

§6. Radioactive Phenomena.

In the present paper it has been attempted to show that the

(1913)) we get in very close agreement

ACCORDING to Rutherford's theory of the structure of atoms, the

§2. Configurations and Stability of the Systems.

Let us consider a system consisting of two positive nuclei of equal

electrons. Denoting this force by , we get from the condition of

The total energy necessary to remove all the charged particles to

For the system in question we have

a table of is given in Part II. p. 32

where has the same signification as in Part II., and

denotes the component, perpendicular to the plane of the

By help of this expression and using the table for given

§3. Systems containing few Electrons. The Hydrogen Molecule.

from (4) we further get

Since , it follows that two hydrogen atoms combine into a

We have assumed in Part I. and Part II. that the frequency of

The value for the frequency of the ultra-violet absorption line in

Let us now consider a slow displacement of the system during which

constant. Putting , we have seen on p. 55 that the radius

Introducing and , we get

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9 How Genetic Tools Can Help Crocodilians’ Management

1.2 Geographic Distribution

1.5 Conservation Status