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PATTERN OF SPECIATION OF FELIDAE

ZIA CZARINA A. GARCIA

A Synthesis Paper Submitted to the Faculty of the Department of the Biological


Sciences, College of Science, Central Luzon State University,
Science city of Muñoz Nueva Ecija, Philippines in
Partial Fulfillment of the Requirements
for the Subject of

EVOLUTIONARY BIOLOGY
(BIOL 3600)

MAY 2021
INTRODUCTION

Morphology of Felidae

Felidae is under the Class Mammalia and under Order Carnivora. Felids showed

variety of body types throughout their worldwide distribution. According to Pallandre,

2019, their body weight ranges from 1 kg (small cats) up to more than 300 kg (bigger

cats, example Panthera tigris). The two optimum body mass of cats are around 5

kilograms for smaller cats and more than 300 kilograms for bigger cats. There is a

striking likeness between all felids. Felids have a shorter rostrum and tooth row than

members of the Canidae family, which improves bite force. The loss or reduction of

cheek teeth is most noticeable in felids, whose dental formula is 3/3, 1/1, 3/2, 1/1 = 30

(Boorer, 1970). The upper premolar is severely reduced in most species, and has been

completely eliminated in Lynx. Carnassials are well developed in felids. They have

secodont cheek teeth that are optimized for shearing. Felid canines are long and conical

in shape, making them perfect for puncturing prey tissue with little force. Felids have big

bullae split by a septum, no alisphenoid canal, and paroccipital processes flattened

against the bullae, in addition to a short rostrum. Felids also have retractable claws and a

vestigial or missing baculum. An elastic section pulls the distal segments of digits back

and up into a sheath in the relaxed position, preventing claws from becoming blunt (

Denis, 1964). Cheetahs are an exception since they are unable to retract their claws and,

while attacking prey, they like to run into them and knock them down, similar to canids

(Grizmek, 2003). On their forefeet, cats have five toes and four on their hindfeet. Their
metapodials are somewhat lengthy but never fused, and they are digitigrade (Walker,

1975).

Males are larger and more muscular than females, exhibiting sexual dimorphism.

Males of some species, such as lions (Panthera leo), may have adornment to attract

possible mates (Boorer, 1970). Feline coats are the longest where temperatures are the

coldest across their range (Feldhammer, et al., 1999). Felids come in a variety of colors,

ranging from black to orange to white, and many species have cryptically colored coats

with rosettes, spots, and stripes to aid in prey detection. In several animals, melanistic

variations (solid black) are widespread, whereas fully white individuals are uncommon.

Within one species, there is a lot of color variety, and infants have a different colour than

adults. Adult cougars (Puma concolor), for example, rarely have spots, although kittens

almost invariably do (Etnyre et al., 2011). The ventral surface of felids is typically pale,

with black or white patterns on the face, tail, and back of the ears. Felids have a variety of

morphological modifications that have enabled them to become the most skilled hunters

in the Carnivora group. They have a digitigrade posture, which results in a fast stride rate

and powerful forelimbs, which aid in the catch and retention of huge prey. Felids are

frequently cryptically colored, which aids in hunting camouflage. Furthermore, most

felids have huge eyes with excellent vision (Kelsey-Wood, 1989). The tapetum lucidum

aids in the intensification of scarce light in nocturnal species (Walker, 1975). Large semi-

rotating ears are also found in many animals. Finally, the felid tongue has a sandpaper-

like feel as a result of posteriorly directed papillae on its dorsal surface, which are

supposed to aid in food retention (Mckenna et al.,2011).


Figure 1. The eight lineages of Family Felidae with descriptions (O’Brien & Johnson,
2007).

Diversity of Felidae

During the Eocene, the first cat-like mammals appeared some 60 million years

ago (MYA), culminating in Barbourofelis fricki, the most specialized of the saber-tooths

(Boorer, 1970). The phylogeny of saber-toothed cats and their forebears (Nimravidae) is,

however, hotly debated, and there is no fossil evidence for these cat-like mammals after
the Miocene (Colby, 1964). Although early ancestors of modern felids had short upper

canines, felid radiations that occurred during the Miocene and Pliocene, such as

Smilodon, appeared to specialize on huge herbivores and possessed enormous, saber-like

upper canines. Machairodontinae (saber-toothed cats) and Felinae (feral cats) were the

two subfamilies of early felids (conical-toothed cats). The three tribes of saber-toothed

cats are separated into many genera (Metailurini, Homotheriini, and Smilodontini).

Conical-toothed cats, both living and extinct, are classified as a single subfamily and

tribe, the Felini, however there is debate over felid categorization at the generic level

(O’Brian, 2001).

Figure 2. The Cat Family Tree according to O’Brien & Johnson, 2007, it contains details
about the ancestors of the 37 species of Felidae.
Native populations of cats can be found all over the world, with the exclusion of

Antarctica, Australia, New Zealand, Madagascar, Japan, and most oceanic islands, and

one species, domestic cats, has been imported practically anywhere humans currently

reside. Despite the fact that certain sources only recognize a few genera, most Felidae

descriptions list 18 genera and 36 species (Lande et al., 2011). With the exception of the

larger cats, the majority of cats are excellent climbers and swimmers. The majority of

felids live alone. Cats are frequently divided into two subgroups: giant cats and tiny cats.

Small cats are those that are unable to roar due to a hardening of the hyoid bone

(Vaughan et al., 2000).

Distribution of Felidae

According to Etnyre, et al., 2011, apart from Australia and Antarctica, felids are

found on every continent of the world.

Figure 3. The migration of cats, according to O’Brien & Johnson, 2007.


On the planet, there are 38 different species of cats. The majority of them are

quite little. The phrase "big cat" refers to any of the five Panthera species: lion (Panthera

leo), tiger (Panthera tigris), jaguar (Panthera onca), leopard (Panthera pardus), and

snow leopard (Panthera pardus) (Panthera uncia). Except for the snow leopard, all of

these big cat species have the ability to roar. A broader definition of Panthera includes

animals such as the cougar (Puma concolor), clouded leopard (Neofelis nebulosa), Sunda

clouded leopard (Neofelis diardi), cheetah (Acinonyx jubatus), and numerous lynx

species. These new species, on the other hand, do not roar.

Figure 4. Geographical distribution of Big Cats except Australia © Vividmaps


Figure 5: Geographic distribution of Felid species © Mattern & McLennan, 2000.
Significance

The evolutionary process by which a new species arise is called speciation. There

are only few studies about how Felid species arise or there are few topics about the

pattern of speciation of Felids. Since Cat is my favorite animal, I am curious about how

their species arise and become adorable as they are now.

REVIEW OF RELATED LITERATURES

According to Mattern and McLennan (2000), the mixture of the data regarding

phylogenetic, distributional, and ecological proved that vicariant speciation has played a

minor role in Felid diversification (approximately 26% of events), whereas sympatric

speciation has been more important than expected on theoretical grounds (approximately

51.8 percent of events), though post speciation dispersal may have blurred the boundaries

between sympatric, parapatric, and peripheral isolate modes. An investigation of Felid

tree ecological changes reveals that resource partitioning patterns in time (activity

patterns), space (preferred habitat type), and food (as indicated by body size) are all

replicated among closely related species. Thus, rapid cat diversification appears to have

been linked to ecological rather than geological separation.

According to Randi et al (2001), the sympatric speciation of cats or crossbreeding

of wild cats with free-ranging domestic cats is thought to jeopardize the genetic integrity

of wildcat populations in Europe, yet diagnostic indicators for determining whether a

wildcat is “pure” or “admixed” have never been defined. Wildcat populations in Eurasia

and Africa are frequently sympatric, and hybridization with domestic cats is a possibility.
Introgression could have been going on for generations, blurring the line between wild

and domestic gene pools. Wildcat populations in Eurasia and Africa are frequently

sympatric, and hybridization with domestic cats is a possibility. Introgression could have

been going on for generations, blurring the line between wild and domestic gene pools.

Cats' behavior and coat colors changed dramatically as a result of domestication.

Behavioral repertoires, on the other hand, are malleable and can quickly adjust to

changing environmental conditions. Natural selection against domestic coat color traits,

on the other hand, might quickly reconstruct wild-type phenotypes in wild-living hybrid

cats. As a result, it's been difficult to create morphological characteristics that can be used

to distinguish pure wildcats under these settings. They concluded that The wildcat is

threatened and declining throughout most of its territory, despite national and

international protection in most European nations, due to habitat destruction, direct

persecution, unintentional death, viral illness transmission, and probable hybridization

with feral cats. To enforce legal protection, it's crucial to improve a set of morphological,

behavioral, and molecular qualities that may be used to identify wildcats, as well as map

the geographic distribution of "pure" wildcat populations that must be preserved as soon

as possible.

Li et al (2015), states that although interspecies hybridization has recently been

identified as a possible regular occurrence in wild animals, the extent to which it shapes

current genomes is yet unknown. To distinguish historical hybridization events from

other processes that cause phylogenetic discordance among different markers, future

researchers need a well-resolved species tree that takes into account all modes of

inheritance and uses large genomic character sets to overcome systematic problems
caused by rapid lineage diversification. Phylogenetic discordance signatures were found

in abundance in the genomes of modern cats, indicating that hybridization was the most

likely source in many cases. X-linked divergence times were significantly reduced across

several large recombination coldspots, which were highly enriched for signatures of

selection-driven post-divergence hybridization between the ancestors of the snow leopard

and lion lineages, according to a comparison of big cat whole-genome sequences. These

findings point to the mosaic origins of current felid genomes, as well as the role of sex

chromosomes and sex-biased dispersal in post-speciation gene flow.

According to Brian et al (2010), the five large cats of the genus Panthera, the lion,

tiger, jaguar, leopard, and snow leopard, as well as the closely related clouded leopard,

separated from the rest of the present Felidae lineage less than 11 million years ago.

There is a considerable difficulty with the precise phylogeny of these critically

endangered large cats. Despite the fact that there have been numerous publications on the

issue, no two molecular investigations have recreated Panthera with the same topology.

Due to the recent and fast spread of pantherines in the Pliocene, individual speciation

episodes happening in less than 1 million years, and likely introgression between lineages

following their divergence, these evolutionary connections remain unexplained. Since

there are lots of queries about the speciation of cats, the authors formulated a supermatrix

and species tree methods to solved the problems. Their findings show a strong consensus

topology supporting lion and leopard monophyly, with jaguar as a sister species, as well

as a sister species relationship between tiger and snow leopard. These findings open up

new options for the study of speciation genomics and the understanding of the historical

events surrounding the lineage's inception.


According to McPete (2014), there are many hypotheses that existed for decades

to explain the origins of cats. Most researchers used to believe that all domestic cats

descended from a common progenitor in ancient Egypt, when cats were tamed around

3,600 years ago. Other experts felt that domestic cats had multiple ancestors, as a result

of domestication occurring at several times and locations, each resulting in a distinct

ancestral breed. The Mediterranean island of Cyprus has the earliest archaeological

evidence of cats and people living together. According to her a research published in

Scientific America was posted online at catoddities.com, stating a 9,500-year-old burial

site on the island of Cyprus was discovered in 2004 by Jean-Denis Vigne of the National

Museum of Natural History in Paris. The skeleton of an 8-month-old cat was discovered

among the many artifacts recovered in the burial, including a chunk of iron oxide, stone

tools, and seashells. The cat was laid to rest in its own tomb, facing westward toward the

setting sun, just like the human remains next to it. Because cats are not native to

Mediterranean islands, the cat buried there was almost certainly transported there by

human immigrants from the Levantine coast (now Turkey, Syria, Lebanon, and Israel).

All domestic cats are descended from Feline silvestris lybica, the Near Eastern wildcat,

according to both archaeological and genetic data, and domestication began in the Middle

East. It's thought that wildcats were drawn to human settlements to hunt mice when mice

invaded human environments in quest of food. Cats were valued by early farmers for

their ability to manage the rodent population. Selective pressure aided in the development

of tameness while preserving rodent hunting abilities. As people travelled around the

globe, cats either followed their human partners or were purposefully brought along.
CONCLUSION

Speciation occurs in many ways; allopatric, sympatric, parapatric, peripatric, and

quantum speciation. I have learned that the pattern of speciation of Felids is allopatric

and sympatric, allopatric in a sense that they interbreed in their own mainland but due to

separation of islands and population a new species arise and sympatric in a sense that

because of evolution they cannot reproduce with each other anymore.

The methods used in the studies are the phylogenetic analysis, which taught me to

be more observant in analyzing a tree; the phylogenetic tree reconstruction; choosing the

characteristics to be used in analyzing the phylogenetic tree, the characteristics are

Karyotypes, Morphology and Molecular sequences which enables researchers to target

the ancestors of the species. The Character Mapping of the species enables researchers to

know the traits of the species. The Modes of speciation, this methods allowed researchers

to identify the speciation of the organism.

The DNA Analysis and Mitochondrial DNA Sequencing and Microsatellite

Genotyping, enables researchers to identify the genetic information of the species and to

be able to know if the organism is wild or domesticated. The Bayesian Clustering method

enables researchers to identify which are the hybrids from domesticated and wild species.

Most of the methods used were focused on the genetic information of the species

since genes are the basic unit of heredity these are the keys of knowing the ancestors of

another species. Studying the molecular level of species will enable researchers how they

interbreed and how does the new species arise.


LITERATURE CITED

Boorer, M. 1970. Wild Cats. New York: Grosset & Dunlap.

Davis, J., R. Anderson. 1971. Parasitic Diseases of Wild Animals. Ames, Iowa: The Iowa
State University Press.

Denis, A. 1964. Cats of the World. Boston: The Riverside Press Cambridge.

Feldhamer, G., L. Drickamer, S. Vessey, J. Merritt. 1999. Mammalogy: Adaptation,


Diversity, and Ecology. Second Edition. Boston, MA: McGraw Hill.

Grzimek, B. 2003. Cats (Felidae). Pp. 369-392 in H Michael, K Devra, G Valerius, M


Melissa, eds. Family: Felidae, Vol. 14, 2 Edition. Farmington Hills, Michigan: The
Gale Group.

Kelsey-Wood, D. 1989. The Atlas of Cats of the World: Domesticated and Wild.
Neptune City, NJ: T.F.H. Publications.

Li G, Davis BW, Eizirik E, Murphy WJ. (2016). Phylogenomic evidence for ancient
hybridization in the genomes of living cats (Felidae). Genome Res. 2016
Jan;26(1):1-11

Mattern, M. Y., & McLennan, D. A. (2000). Phylogeny and speciation of


felids. Cladistics, 16(2), 232-253.

O'Brian, S. 2001. Cat Family. Pp. 8-39 in D Macdonald, ed. The Encyclopedia of
Mammals. Oxford, UK: Andromeda Oxford Limited.

O’Brien, S. J., & Johnson, W. E. (2007). The evolution of cats. Scientific


American, 297(1), 68-75.

Randi, E., Pierpaoli, M., Beaumont, M., Ragni, B., Sforzi, A., (2001). Genetic
Identification of Wild and Domestic Cats (Felis silvestris) and Their Hybrids Using
Bayesian Clustering Methods, Molecular Biology and Evolution, Volume 18, Issue
9, September 2001, Pages 1679–1693

Vaughan, T., J. Ryan, N. Czaplewski. 2000. Mammalogy. Fourth Edition. Philadelphia,


PA: Brooks/Cole-Thomson Learning.

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