Rana+(Lithobates)+Catesbeiana

American Bullfrog  **//Rana// (//Lithobates//) //catesbeiana//** **(Shaw, 1802)**

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 * TOP 10 BULLFROG FACTS **
 * 1) Bullfrogs can leap up to 10x body length (2 meters)
 * 2) Weigh up to 1 kg
 * 3) Adults can grow up to 18 centimeters in length (excludes legs)
 * 4) Longest lifespan : 16 years in captivity, 10 years in the wild
 * 5) Calls excessively when thunderstorm approaches
 * 6)  Food delicacy : J uicy and meaty hind legs
 * 7) Asian dessert : Dried fatty tissues of frog's ovaries have anti-aging properties (Hasma or //Oviductus ranae//)
 * 8) Major invasive species originating from North America
 * 9)  Resistant to lethal amphibian fungal infection  (chytridiomycosis)
 * 10) Efficient carrier of 'chytridiomycosis-causing fungus' ( // Batrachochytrium dendrobatidis // )

=** Introduction **=
 * Largest frog species in Singapore
 * Aquatic frog, belonging to the family 'true frogs' (Ranidae)
 * Bulk of //Rana// (//Lithobates//) //catesbeiana// population is with the commercial breeding company (Jurong frog farm)
 * Reared and exported the frogs as food
 * Increase sightings of //Rana// (//Lithobates//) //catesbeiana// in Singapore's water catchment areas.

**Etymology **
The genus name '//Rana//' is a Latinized word meaning frog, is a proposed genus name by George Shaw in 1802. However in 2006, John S. Frost replaced the genus to '//Lithobates//', which is a combination of two Greek words 'λίθος' and 'βαίνω'. The word 'λίθος' stands for gemstone, while the word 'βαίνω' stands for walk. The species name '//catesbeiana//' is to honor Mark Catesby who was a famous English naturalist who first discovered the bullfrog. The later revision by Green (2007) of the species name '//catesbeianus//' is a new Latin patronym for Mark Catesby.

=** Physical Traits **=

Adults


__Overall body __
>
 * 1) Smooth and non-warty skin
 * 2) Absence of dorso-lateral ridges
 * 3) Adult dorsal coloration range from greenish, to yellowish or olive-blackish.
 * 4) Ventral coloration is usually whitish or yellowish accompanied with dark mottling patches.
 * 5) Head usually a lighter shade of green, with its legs being darkly blotched or banded
 * 6) General coloration varies widely according to habitat/location

__Limbs __

 * 1) Long and muscular hind legs
 * 2) Well developed webbing in between the toes of hind feet
 * 3) Anterior limbs are much shorter than the hind legs and are used to prop the frog's posture up tall.
 * 4) No webbing is seen between the fingers of the anterior limbs
 * 5) Both the tips of the frog's fingers and toes are blunt

__Head __
>
 * 1) Broad snout
 * 2) Small nostrils are situated near the tip of the snout
 * 3) Teeth in its upper jaw
 * 4) Vocal openings at the corner of the mouth Large eardrums (tympanums)
 * 5) Supra-tympanic fold just above the tympanum, starting from the eye and past the eardrum and finally extending towards its anterior limbs.
 * 6) Prominent eyes that are situated above the head unlike the tadpoles which have eyes on the sides of their heads
 * 7) Almond-shaped horizontal pupils and brown irises.

Diagnosis
 General diagnostic traits for comparison with ** other frog species ** :

media type="youtube" key="mtAdhpTKmgg" width="616" height="346" align="center"
 * 1) True frog appearance
 * 2) Smooth skin
 * 3) Webbed feet
 * 4) Long hind legs that represent about half its body length and 40% of its body weight.
 * 5) Deep and resonant mating call that resembles the moo of the cow (hence its vernacular name, the American bullfrog.)

[|Rana catesbeiana calling in a mating pond]  Distinct traits for comparison with ** closely-related ranid frog species ** : This distinctly identifies the American bullfrog (//Rana// (//Lithobates//) //catesbeiana//), which is often mistaken for the river frog (//Rana hecksheri//), pig frog (//Rana grylio//) and green frog (//Rana clamitans//) since the geographical ranges of these three species overlap with //Rana// (//Lithobates//) //catesbeiana//'s native range.
 * 1) Large tympanums
 * 2) Prominent supra-tympanic folds
 * 3) Lack of dorso-lateral ridges

 * The tympanums are the circular eardrums that are situated right behind each eye of the frog.

Sexual Dimorphism
 __ **Summary of sexual dimo****rphism** __
 * ** Trait of comparison ** || ** Male Bullfrog ** || ** Female Bullfrog ** ||
 * Time to reach sexual maturity || 1 to 2 years || 2 to 3 years ||
 * Average snout to vent length || 152 mm || 162 mm ||
 * Tympanum size || Twice the size of eye || Same size as eye ||
 * Ventral throat coloration || Yellowish || Whitish ||
 * Pigmented nuptial pads || Present || Absent ||
 * Additional notes || - || Retain some juvenile color and morphology ||

**Juvenile Adults **

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 20pt;">[[image:Rana catesbeiana juvenile black .jpg align="right" caption="<Photo taken by Ip Yin Cheong Aden>"]]
<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14pt;">Juvenile adults or frog-lets are the intermediate transition phase of the metamorphosed tadpoles that have re-absorbed their gills and finned tails. They are miniatures of adult bullfrogs.

> > <span style="font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> * The parietal eye is homologous to a third eye and is capable of sensing changes in light intensity related to day and nightfall, which is also use for synchronization of its biological clock ([|circadian rhythmicity]).
 * Average 4 cm from snout to vent
 * Do not have sexually dimorphic characteristics yet
 * Orange irises in their eyes
 * Possess a grey oval shaped skin area between their eyes which is called the parietal eye

**<span style="font-family: Arial,Helvetica,sans-serif; font-size: 20pt;">Larvae (Tadpoles) **
<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> <span style="font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> Tadpoles pump water through their gills by movements of the floor of their mouths, trapping bacteria, protozoans and other small particles on mucus in a filtration organ in their pharynges. <span style="font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> As they grow, they begin to ingest larger particles and use their teeth for rasping. <span style="font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> Growth of hind legs are usually observed before the anterior limbs bud out while the tadpole metamorphoses into a frog-let
 * Tadpoles are also very large by frog standards, 8cm to 15cm
 * Long, muscular tail with broad dorsal and ventral fins
 * Downward-facing mouths
 * Dorsal coloration is brown to light olive with small black spots scattered across the head and upper body.
 * Ventral side is usually whitish to yellowish in color.
 * Initially have three pairs of external gills, and several rows of labial teeth
 * Unlike the adult frogs, tadpoles have their eyes at the sides of their heads with orange irises

==**<span style="font-family: Arial,Helvetica,sans-serif; font-size: 20pt;">Eggs **==
 * Very small
 * Dorsal black color
 * Slightly lighter or whitish ventral underside
 * Each egg is surrounded by a jelly capsule
 * Additional jelly that creates a loose cohesion to the entire mass of eggs
 * About 10,000 to 20,000 eggs are usually laid near the water surface in one brood

=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 30pt;">Biology = <span style="font-family: Arial,Helvetica,sans-serif; font-size: 20pt;">**General behavior**
 * ** Trait ** || ** Adults ** || ** Larvae ** ||
 * Activity || Principally nocturnal, though observed to be active throughout the day || Principally nocturnal, though observed to be active throughout the day ||
 * Residing environment || Semi aquatic and terrestrial || Fully aquatic ||
 * Feeding type || Carnivorous || Herbivorous ||
 * Muscular hind legs / tail || Used to leap for prey, grasp females or simply for escaping || Used to support and maneuver its huge body in water ||

**<span style="font-family: Arial,Helvetica,sans-serif; font-size: 20pt;">Types of defensive behavior **
<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14pt;">Mature male //Rana// (//Lithobates//) //catesbeiana// are extremely territorial animals. The male frogs do not hesitate in attacking a competitor or intruder of their turf, with rapid kicking, leaping and grappling.
 * ==== Grappling for Defense ====

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14pt;">//Rana// (//Lithobates//) //catesbeiana// have large mouths with a row of teeth in the upper jaw, rendering it capable of inflicting painful bites onto unsuspecting targets. The small teeth on its upper jaw are effective tools to help the bullfrogs to grab it prey as well.
 * ==== Biting in Defense ====

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14pt;">//Rana// (//Lithobates//) //catesbeiana// often uses defensive vocalization for self-protection. Their vocal defensive behavior acts to ward off danger to both conspecifics and predators. The defensive call,also known as the distress call, is rapid, unpleasant sounding and piercing for its predator.
 * Vocalization for Defense

media type="youtube" key="Q9_kckU5O50" width="560" height="315" align="center"

[|Rana catesbeiana distress call <at 0.16 of video>]

**<span style="font-family: Arial,Helvetica,sans-serif; font-size: 20pt;">Mating Behavior **
<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> * Males only clasp females in the amphiplexus position, after they have signaled their willingness for mating. This debunks the prior belief that a male frog will opportunistically clasp any nearby female to mate regardless of her consent.
 * Males are extremely territorial
 * Occupy sites that are usually spaced 3 to 6 meters from the next frog and call loudly
 * Males make different types of calls, such as territorial calls to warn other males
 * Males make advertisement calls to attract females

=<span style="color: #170202; font-family: Arial,Helvetica,sans-serif; font-size: 30pt;">Habitat and Reproductive cycle = <span style="color: #150101; font-family: Arial,Helvetica,sans-serif; font-size: 15pt;">//Rana// (//Lithobates//) //catesbeiana// can thrive in a wide range of environmental conditions since they are a species of frogs that easily adapt to the environment. They are found in both temperate where they hibernate over winter, as well as tropical regions where temperatures can go up to 40 °C.

<span style="color: #150101; font-family: Arial,Helvetica,sans-serif; font-size: 15pt;"> The duration of the breeding season is influenced by the amount of sunlight, temperature, atmospheric pressure and humidity. The closer //Rana// (//Lithobates//) //catesbeiana// is to the equator, the longer its reproductive period through the year. It has been observed that //Rana// (//Lithobates//) //catesbeiana// breeds all year round in Singapore, but only have a three-month breeding season in North America.

<span style="color: #150101; font-family: Arial,Helvetica,sans-serif; font-size: 15pt;"> Once the eggs are laid, the development of embryos is determined by temperature, whereby larvae will hatch within 48 hours in warmer conditions above 26 °C. Egg masses are laid near the water surface for facilitation of oxygen diffusion into eggs. Larvae are about 10mm long, attach to smooth surfaces and consume their yolk sac within 72 hours. After the yolk sac has been emptied, the larvae become free swimming and filter feeding tadpoles.

<span style="color: #b22c46; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;">

__ Larvae __
<span style="color: #120709; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> The occurrence of metamorphosis in tadpoles depends on two environmental factors which also act as the forces for natural selection:
 * Hatch in about 4 days to be fully aquatic and gilled tadpoles
 * Tadpole stage may persist for up to 2 years
 * The longer the time taken for a tadpole to metamorphose, the larger the adult would be

<span style="color: #a60f21; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> *Decreased competitive interactions and more favorable environmental conditions (food, high temperature and clean water), would accelerate the process of metamorphosis from tadpoles to adults.
 * 1) Competitive interaction; includes predator, inter and intraspecific interactions
 * 2) Environmental conditions; food availability, temperature, water quality

__<span style="font-family: Arial,Helvetica,sans-serif; font-size: 16pt;">Adults __
media type="youtube" key="UHpTH0ZWBJI" width="560" height="315" align="center"
 * After one or two years, tadpoles metamorphose into adults
 * Reproduction occurs in permanent water bodies (In Singapore since its warm all year round, breeding season is all year, especially after episodes of heavy rainfall)
 * Males attract females by calling (a deep, hoarse "ba-rum" or "jug-o-rum")

[|Rana (Lithobates) catesbeiana calling and mating]
 * Mating occurs in water; males clasp females and hold them in amphiplexus
 * Females release eggs while in amphiplexus; eggs are surrounded by a gelatinous sheet that adheres to vegetation
 * Fertilization is external
 * Average lifespan of adults is about 6 years

=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 30pt;">Diet = media type="youtube" key="vK6HmAxmjM4" width="560" height="315" align="center"
 * Adults usually ambush its unsuspecting prey by sitting quietly
 * As the prey passes by, it lunges forward with a wide open mouth, powered by its powerful hind legs.
 * The bullfrog eats and swallows practically anything that can fit into their mouths.
 * They have a wide range of prey, from insects to mice, fish, birds, herpetofauna (snakes and turtles) and even its own kind (cannibalism).

[|American bullfrogs eat anything that fits into its mouth!] media type="youtube" key="kVfZWA30dFQ" width="624" height="354" align="center" > >
 * Bullfrog tadpoles mostly graze on aquatic plants, or eat suspended matter, organic debris, algae, plant tissue, and small aquatic invertebrates
 * During metamorphosis, diet gradually shift towards ingesting animal protein
 * Fully carnivorous after tail resorption in the froglets
 * [|A day in the life of a bullfrog tadpole]

=** Distribution **=

__**<span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Global **__
**<span style="color: #150101; font-family: Arial,Helvetica,sans-serif; font-size: 15pt;"> *Adapted from [|CABI]; direct permission not obtained, but [|within limits of Fair Use] **

__**<span style="font-family: Arial,Helvetica,sans-serif; font-size: 16pt;">Singapore **__
[|Click here to access interactive Google Map for local American Bullfrog Sightings.] <span style="color: #150101; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> *The objective of this map is to allow the general public to mark out the places where they have spotted the American bullfrogs in Singapore. This would aid in monitoring and assessment of the spread of this invasive frog species with respect to local ecosystems, native anuran species richness and ecological balance.

<span style="color: #150101; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> The American bullfrog is locally found in the water catchment areas as well as drains, canals and park connectors

<span style="color: #150101; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> Sighted in: >
 * 1) Macritchie Central Catchment (Most sighted)
 * 2) Seletar reservoir
 * 3) Peirce reservoir
 * 4) Pandan reservoir
 * 5) Punggol reservoir
 * 6) Marina reservoir
 * 7) Kranji reservoir
 * 8) Jurong lake
 * 9) Bedok reservoir

=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 30pt;">Invasive Species Implications = > >
 * Imported into Singapore since American bullfrog legs are a profitable and marketable consumable food commodity
 * Easily cultivated but also difficult to contain as they are very prone to escaping (leaps up to 2 meters)
 * Used in aquatic pet trade as baits or food for the Arowana fish (F. Osteoglossidae)
 * Accidental release or escapes result in thriving populations with thousands of bullfrogs within a year of colonization arising from just one successful spawning event.
 * They are highly adaptable to new environments and are large, opportunistic predators that out-competes native anurans.
 * Larvae rapidly eat algae especially in nutrient poor ponds, monopolizing primary production and upset aquatic community structure
 * The American Bullfrog is resistant to Chytridiomycosis and hence is a well known vector for transmitting this deadly amphibian disease.
 * The chytrid fungus, //Batarachochytrium dendrobatidis//, causes the disease that has wiped out many anuran species thus far
 * Listed as the top 100 most devastating invasive species by the Global Invasive Species Database
 * DO NOT release your pet American bullfrogs, prevent the bullfrog baits and commercially farmed frogs from escaping into our local ecosystem!

=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 30pt;">Taxonomy =

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 20pt;">Taxonavigation

 * Domain: Eukaryota
 * Kingdom: Metazoa
 * Phylum: Chordata
 * Subphylum: Vertebrata
 * Class: Amphibia
 * Order: Anura
 * Family: Ranidae
 * Genus: //Rana// (Shaw, 1802)
 * Sub-Genus: //Lithobates// (Frost et al., 2006)
 * Species: //Rana catesbeiana// (Shaw, 1802)
 * Species (Revised and Preferred): //Lithobates// //catesbeianus// (Green, 2007)
 * Species (CABI (2015) advisory): //Rana// (//Lithobates//) //catesbeiana//

**<span style="color: #110104; font-family: Arial,Helvetica,sans-serif; font-size: 25pt;">Names **

 * Binomial: //Rana// //catesbeiana// (Shaw, 1802)
 * Vernacular: **American bullfrog**, Common bullfrog, Edible bullfrog, Eastern bullfrog, Jug-'O'-rum, Bloody nouns
 * Synonyms:
 * //Rana catesbeiana//, **//Rana// (//Lithobates//) //catesbeiana//** (Shaw, 1802)
 * //Rana// (//Rana//) //catesbeiana// (Boulenger, 1920; Dubois, 1978)
 * //Rana catesbyana// (Smith, 1978)
 * //Rana// (//Aquarana//) //catesbeiana// (Dubois, 1992)
 * //Rana// (//Novirana//, //Aquarana//) //catesbeiana// (Hillis and Wilcox, 2005)
 * //Lithobates catesbeianus// (Frost //et al//., 2006)
 * //Lithobates// (//Aquarana//) //catesbeianus// (Dubois, 2006)

<span style="color: #254daf; font-family: Arial,Helvetica,sans-serif; font-size: 20pt;">Which name do we use?
<span style="color: #140a0c; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;">This frog species was shifted by into the genus //Lithobates// based on extensive sequence data, which is highly debatable because if //Lithobates catesbeuanus// was used, a lot of literature on //Rana catesbeiana// prior to this name change would be lost. Hence, Hillis and Wilcox (2005) proposed //Lithobates// as a subgenus, whereby it is used in a different sense for smaller group of species. Although notable organizations such as the International Union for Conservation of Nature (IUCN), Encyclopedia of Life (EOL) and Animals and Plants of Singapore (APS, LKCNHM) use either //Lithobates catesbeianus// or //Rana catesbeiana//. The Centre for Agricultural and Biosciences International (CABI) also advised to use //Lithobates// as a subgenus and this frog's name could be expressed as //Rana// (//Lithobates//) //catesbeiana//. This would thus maintain and preserve the long historical build up of information on phylogeny and taxonomy of //Rana// (//Lithobates//) //catesbeiana.//

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 20pt;">Type information
<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;">George Shaw (1802) is an English naturalist who gave this bullfrog its original name //Rana catesbeiana//. He gave the type locality (//R//. c//atesbeiana//) as “North America”, although that was later restricted to “vicinity of South Carolina”, Charleston by Schmidt (1953). A type specimen is not known to exist. The collector and date of collection are unknown.

<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> **<span style="color: #b22c46; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> *Present holotype locality status: Unknown **

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 20pt;">Original description


<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 9pt;"> Specimen illustrated by Shaw, 1802, Gen. Zool., 3(1): 106, pl. 33 (Permission pending, University of Michigan Library Administration)

<span style="color: #0b0103; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> Size very large (maximum SVL 203mm in females). Dorsum green to almost black, or with net-like pattern of gray or brown on green ground; venter whitish, mottled gray (at times), and with a yellowish wash, especially on the throat of males; no dorso-lateral ridges; feet with tip of fourth toe extending beyond web; digital disks absent.

<span style="color: #0b0103; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> **<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 20pt;"> More recent illustrations **<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 16pt;">

<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 12px;"> (Permission pending, University of Michigan Library Administration)

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 20pt;">Phylogenetics
<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;">A phylogenetic study of Amphibia by Pyron and Weins (2011) has conducted a maximum likelihood analysis on Amphibia phylogeny based on 12,871 base-pairs of sequence data from the concatenation of 12 genes (9 nuclear, 3 mitochondrial), covering both higher and lower taxonomic levels (up to 2800 amphibian species).

<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 12pt;"> 9 nuclear genes : > <span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 12pt;">3 mitochondrial genes : > <span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> The placement of //Rana// (//Lithobates//) //catesbeiana// on the amphibia phylogenetic tree is represented in the 3 figures below. The figures are presented in a chronological order that goes down the Taxonomic hierarchy. Starting from the Class, Order, Superfamily and Family [Fig. 1]; proceeding on to Family and Genus level [Fig. 2]; and lastly down to Genus and Species level [Fig. 3].
 * 1) C-X-C chemokine receptor type 4 (CXCR4)
 * 2) Histone 3a (H3A)
 * 3) Sodium–calcium exchanger (NCX1)
 * 4) Pro-opiomelanocortin (POMC)
 * 5) Recombination-activating gene 1 (RAG1)
 * 6) Rhodopsin (RHOD)
 * 7) Seventh-in-absentia (SIA)
 * 8) Solute-carrier family 8 (SLC8A3)
 * 9) Tyrosinase (TYR)
 * 1) Cytochrome b (cyt-b)
 * 2) Mitochondrial ribosomal subunit 16S
 * 3) Mitochondrial ribosomal subunit 12S

<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> ** Bootstrap values for maximum likelihood have been indicated on the tree branching nodes for all 3 figures below. (Adapted from [|Science Direct]; direct permission not obtained, but within limits of Fair Use) **

<span style="color: #b22c46; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> <span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> ** Figure 1. Skeletal representation of the maximum likelihood analysis of the concatenated 12 genes of the Orders, Superfamilies and Families in the Class Amphibia. ** <span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 10pt;"> Moving down the taxonomic hierarchy, the right vertical lines denote the 3 Orders, while the node bases denote the superfamily and line tips represent family. //Rana catesbeiana// is in the Order Anura, Superfamily Ranoidea and Family Ranidae as boxed in red. Bootstrap values are indicated at the branch nodes.

<span style="color: #b22c46; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;">

<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> ** Figure 2. Family Ranidae phylogeny's maximum likelihood estimate based on concatenated sequencing data of 12 genes (expanded from [Fig. 1]) ** <span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"><span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"><span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 10pt;">  //<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 10pt; line-height: 1.5;">Rana catesbeiana //<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 10pt; line-height: 1.5;"> (American bullfrog) isolated in red is seen to be in the same clade as 7 other Ranoid frog species  <span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 10pt; line-height: 1.5;">. This includes the 3 closely related true frogs (annotated with red arrows) namely the R. clamitans (Green frog), R. grylio (Pig frog) and R. heckscheri (River frog), which share the same native geographical range and often mis-identified as //<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 10pt; line-height: 1.5;">R. catesbeiana //<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 10pt; line-height: 1.5;"> by para-taxonomists <span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 10pt; line-height: 1.5;">. <span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 13.3333px;">Bootstrap values are indicated at the branch nodes.

<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 10pt;"> [|*Compare misidentified Ranoid true frogs]

**<span style="font-family: Arial,Helvetica,sans-serif; font-size: 20pt;">Gene sequences **
<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;">The study by Pyron and Wiens (2011) did not utilize the mainstream barcoding genes like mitochondrial gene (COI) or nuclear gene (18s rDNA) for taxonomic delineation. <span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> However, most barcoding data for species delineation published on GenBank commonly involves COI or 18S rDNA. Hence, the following mainstream barcode data has been provided to serve as a linking platform to the respective databases for researchers working on barcoding projects (e.g. eDNA water testing studies that involve //Rana// (//Lithobates//) //catesbeiana//.

<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> The illustrative barcode represents a fragment of DNA sequences, with each base uniquely colored for a rough visual identification of the nucleotide composition makeup of the gene.

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 * ===**<span style="color: #070002; font-family: Arial,Helvetica,sans-serif; font-size: 120%;"> Barcode Color Legend **===
 * **Nucleotide** || **Color Code** ||
 * Adenine (A) || Green ||
 * Thymine (T) || Red ||
 * Guanine (G) || Black ||
 * Cytosine (C) || Blue ||
 * === Mitochondrial gene: Cytochrome C Oxidase Subunit 1 (COI) ===

<span style="color: #120306; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> Illustrative COI barcode (646bp) for //Rana// //catesbeiana// obtained from [|BOLD systems] that was deposited in NCBI GenBank. COI sequence file (.fasta) can be obtained from [|GenBank].

<span style="color: #b22c46; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> <span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> Illustrative 18S rDNA barcode (581bp) for //Rana// //catesbeiana// obtained from [|BOLD systems.] <span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 14pt;"> 18S sequences can be obtained from [|BOLD systems].
 * === Nuclear gene: 18S Ribosomal DNA ===

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<span style="color: #240b0b; font-family: Arial,Helvetica,sans-serif; font-size: 120%;">** References **