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© 2001 Peter Brazaitis11 April [email protected]

A GUIDE TO THE IDENTIFICATION OF THE LIVINGSPECIES OF CROCODILIANS

by

Peter BrazaitisResearch Associate

Science Resource CenterThe Wildlife Conservation Society

Prepared for the inaugural class of the AZA Crocodilian Management SchoolSt. Augustine Alligator Farm, St. Augustine, Florida

5 May to 12 May, 2001

This document may not be copied in any form, or distributed in any form, in whole or in part, without written permission of the author.For distribution to AZA member instructors and registered students only.

Copy for: Wildlife Conservation Society, Science Resource Center 5 June 2008



©2001 Peter Brazaitis 2

TABLE OF CONTENTS

INTRODUCTION ------------------------------------------------------------------------------------------3

CROCODILIAN SPECIES ------------------------------------------------------------------------------5

BODY SCALATION --------------------------------------------------------------------------------------6

DEFINING TERMS----------------------------------------------------------------------------------------7

USE OF KEYS----------------------------------------------------------------------------------------------9

KEY TO FAMILIES OF LIVING CROCODILIANS ----------------------------------------------10

KEY TO THE SPECIES Of LIVING CROCODILIANS -----------------------------------------10

KEY TO THE SPECIES OF THE GENUS Caiman --------------------------------------------- 18

SPECIES ACCOUNTS FOR LIVING CROCODILIANS----------------------------------------23

Family: Alligatoridae -----------------------------------------------------------------------------------23

Family: Crocodylidae ----------------------------------------------------------------------------------42

Family: Gavialidae --------------------------------------------------------------------------------------66

ACKNOWLEDGMENTS --------------------------------------------------------------------------------70

REFERENCES --------------------------------------------------------------------------------------------71




INTRODUCTION

The identification of crocodilians has historically been based largely onmorphological characteristics derived from the examination of skulls, skins, andpreserved juvenile specimens. Molecular genetics technologies have since added a

new powerful tool, which has immense value in its application for the conservation,management, and propagation of endangered and threatened species. Today, we areno longer simply concerned with the accurate labeling of species for educationalpurposes, but must concern ourselves with attempting to preserve genetic diversity untoperpetuity among populations, establishing genealogical lines, and selecting individualanimals for propagation which best exemplify the species. Nonetheless, the simpleidentification of species and individuals, based on their morphological characteristics,remains paramount to the development of conservation and captive managementprograms.

Today, more than ever, the genetic integrity of crocodilian species and their wild

and captive populations are at risk. Wild populations, overall, continue to declinearound the world. Critical habitats continue to be lost to human encroachment,commercial development, and industrial pollution. The cost of preserving human healthand quality of life continue to demand increasing precedence over wildlife conservationand habitat protection.

Perhaps a new and greater threat lies in the very sustainable use initiatives thatare meant to relieve the abusive consumption of endangered and threatened species.Conservation research and funding is all too often funneled toward developingcommercial trade and a profitable return on investments, as a priority over theprotection of wild populations and their habitats. The greatest populations for manyspecies currently resides on commercial farms and ranches, where the primary interestis in producing the greatest number of individuals, at the least expense, in the shortestperiod of time, with the best quality of esthetically useable skin for the exotic leathertrade. Not in maintaining the genetic integrity of the species or population. Non-nativecrocodilian species are increasingly being commercially established in captive breedingpopulations within the habitats of native species, because of the greater value of oneskin type over another in commercial trade. Their inadvertent introduction or release,into the wild population at large, may be simply predicated upon the economics ofcurrent market values and investment profits. The random production of hybridhatchlings for the pet trade, in unmanaged common ponds is now an increasingproblem.

Rapidly advancing DNA technology continues to offer new insights intocrocodilian relationships and ancestry. Previous checklists may have relied onspeciation based only on the limited osteological interpretations of early investigators.This present work does not assume a cavalier approach to the introduction and use of“new species” classifications. In this time of intensive national and international wildlifetrade, regulation, and legal ramifications involving illegal trade, the re-classification ofcommercially valuable species for political or legal purposes may be particularly




relevant. On one hand, there is considerable incentive to divide named protectedspecies into “new species,” or populations, which by name, might temporarily falloutside of existing wildlife regulation. Likewise, it may also be legally expeditious, forsome, to combine limited populations of endangered and non-endangered speciestogether, under a single broad taxon, thus legally making available a broader base of

populations available for utilization.

Beyond scientific description, several additional tests may be appropriate indetermining the validity of new taxa.1. Are the core wild populations represented by the taxon reproductively isolated by

ecological, geographical, geological, or other biological barriers? In other words, isthe genetic integrity of the named taxa populations insured in the wild?

2. How morphologically different is the taxon from other species?3. Does DNA analysis support the morphological and zoogeographical distinctness of

the taxon.?4. Are there designated type specimens collected from vouched localities, and

deposited in a scientific institution, available for future reference in perpetuity?

The inclusion of taxa in this work was a decision made by the author, based oncommon inclusion in current literature, the morphological and genetic distinctiveness ofthe species, and an assessment of the current literature in evaluating the distribution ofthe species. Isolated island forms may be included as a sub-species, races ofCrocodylus palustris, for example; while certain populations of a broad ranging species,such as Crocodylus porosus, may not be included.

This paper is directed at providing a tool, which may be useful in the speciesidentification of living crocodilians. It is based on an assessment of morphologicalcharacteristics which typify each species as a whole, the literature, the examination ofthousands of living animals in zoo’s and in the field, preserved materials in museums,universities, and other collections. More than 800 comprised that database. Not everyindividual animal will display typical species characteristics. Captive produced animalsare more difficult to identify than animals from wild origins, because of artificiallyproduced anomalies that may distort some physical characteristics. However, theprimary characteristics displayed should fall within the range of parameters for thatspecies. Given the increasing incidence of hybrid individuals finding their way intocaptive collections, it may be necessary to seek confirmation of species identity by DNAanalysis. Certainly, those individuals who significantly diverge from representativespecies or population characteristics should not be considered for reproduction or re-introduction programs for endangered or threatened species.




CROCODILIAN SPECIES

The present work is based on the following taxa:

Family Species Common name

Alligatoridae Alligator mississippiensis American alligator Alligator sinensis Chinese alligatorCaiman crocodilus crocodilus common caiman

Caiman c. apaporiensis Rio Apaporis caiman

Caiman c. chiapasius Central American caiman

Caiman c. fuscus dusky or brown caiman

Caiman yacare Yacare caiman

Caiman latirostris broad-snouted caiman

Melanosuchus niger black caiman

Paleosuchus palpebrosus dwarf caiman

Paleosuchus trigonatus smooth- fronted caiman

Crocodylidae Crocodylus acutus American crocodile

Crocodylus cataphractus African slender-snouted crocodile

Crocodylus intermedius Orinoco crocodile

Crocodylus johnsoni Johnstone’s crocodile

Crocodylus mindorensis Philippine crocodile

Crocodylus moreletii Morelet’s crocodile

Crocodylus niloticus Nile crocodile

Crocodylus novaeguineae New Guinea crocodile

Crocodylus palustris palustris mugger, or marsh crocodile

Crocodylus palustris kimbula Ceylon marsh crocodile

Crocodylus porosus saltwater, estuarine,or Indo-pacific crocodile

Crocodylus rhombifer Cuban crocodile

Crocodylus siamensis Siamese crocodile

Osteolaemus tetraspis tetraspis West African dwarf crocodile

Osteolaemus. t. osborni Congo dwarf crocodile

Gavialidae Gavialis gangeticus Indian Gharial

Tomistoma schlegelii Malayan false gavial




BODY SCALATION

Fig. 1. Crocodilian body scalation.

Dorsal scalesDouble caudal

whorls

Single caudal

verticils

Flanks

Lateral tail region

Ventral collar

Ventral scale rows,

collar to vent.

Anal vent

Ventral tail region

(sub-caudal)

Post-occipital scalesNuchal scales

Post-nuchal scales

Sub-caudal

inclusions

Lateral tail

inclusions




DEFINING TERMS

General:Content: Any sub-species or named populations which comprise the taxon.Size: General information on the sizes attained by the species.

Coloration: Color and pattern.Conformation: Information on the shape and form of the species.Scalation: The numerical range and description various scale groups.Dentition: The dental formula, form, arrangement, of the teeth and jaws. It is important

to note that when defining “tooth in an external notch” as opposed to “toothreceived in an internal socket”, care must be taken to avoid mistaken anomaliescaused by dietary deficiencies often found in captive raised animals. The teethof healthy wild crocodilians grow in a vertical plane. That is, straight up anddown, the lower teeth fitting uniformly between the upper teeth when the jaws areclosed, often forming a perfect “zig-zag” pattern. Teeth that protrude outwardfrom the jaw, at the front and sides; jaws that grow as an under-bite or over-bite;

missing teeth that are not replaced; tooth sockets that appear laterally rotated, allsuggest captive anomalies. The first mandibular teeth and the fourth mandibularteeth often pierce, and protrude through the upper jaw, in healthy wild Caiman,when the jaws are closed; and are not visible along the jaw line when the jaw isclosed. Dental anomalies in Caiman are often manifest by teeth that extendradically outward, particularly the first and fourth mandibular teeth. Although aproper socket exists to receive the tooth, the tooth has rotated out of thereceiving socket, giving an erroneous impression of having an “exposed fourthtooth”. The important factor is to look for the presence of a distinct externalnotch on the side of the jaw, which is absent in alligatorids, rather than simplyrelying on a view of the tooth itself.

Counts are given for each of the jaw, and are not total mandibular or maxillarycounts. Example: 5+13-15 indicates there are 5 pre-maxillary teeth, followed by13 to 15 maxillary teeth on each side of the upper jaw. 19-20 below the barindicates there are 19 to 20 teeth on each side of the mandible or lower jaw.Doubling the counts would give total tooth counts for the upper and lower jaws.

Similar Species: Those species in which morphological variation might lead to amisapplication of the key; species which may be confounded with anotherspecies through similarity; alternate species to review when the animal underexamination does not entirely conform to the key results.

Distribution: The general range of regions where the species occurs. It should beremembered that crocodilians do not inhabit regions defined by politicalboundaries, but rather the river systems and wetlands within political regions.

Comments: Important information about the classification or history of the species.References: Important literature.




Scalation:Neck regions:Post-occipital scales: One or several rows of scales located immediately behind the

head at the base of the skull. Types include a single row of conspicuouslyenlarged, highly keeled scales, arranged in two or three equally divided pairs;

several rows of slightly enlarged scales keeled scales; a single pair of onlyslightly enlarged scales; many bead-like scales, without any enlargement.Nuchal scales: Large, heavily boned and closely connected scales, located on the top of

the neck, immediately following the post-occipital scales. Types of nuchalscalation include an oval cluster of usually six scales, a square of four scales, aseries of parallel paired scales that become continuous with the dorsal scales

Post-nuchal scales: Usually two relatively small scales located between the nuchalscales and the first row of four to six dorsal scales. Clearly evident inCrocodylus and Osteolaemus but often difficult to identify from the last nuchalscale rows in alligatorids, Tomistoma, and Gavialis.

Body trunk regions:Dorsal scales: The somewhat keeled scales covering the back of crocodilians. Dorsalscales usually begin with a series of four to six scales, may increase to 10scales per row, reducing to as few as two scales in a row over the hind limbs(lumbar area). Dorsal rows are counted from the first row of scales following thepost-nuchal scales, and ending with the last row of scales that coincide with theposterior base of the hind legs. Nearly all crocodilians possess 17 to 18 dorsalscale rows.

Flanks: The regions on each side of the body between the front and hind limbs, thedorsal scales and the ventral scales. Flank scales consist of conspicuouslyenlarged, longitudinal rows of scales, or a series of individual scales. Theinterspaces of skin between the enlarged flank scales may be composed of smallscales or soft skin. Flank scale rows are counted at mid-body, beginning with thefirst row of enlarged scales below the dorsal scales, and ending with the last rowof enlarged scales bordering the ventral or belly scales.

Ventral collar: A series of often very enlarged scales that form a “mantel”, extending from the anterior shoulder of one front leg, across the base of the neck, to theanterior shoulder of the other front leg. In some species, these scales are notnoticeably enlarged, although the mantel is present.

Ventral scale rows: The transverse rows of belly scales, beginning with the first row ofscales after the ventral collar, and ending with the anterior most point of the analvent. Ventral scale counts are made by counting each row from collar to vent. Inmany cases, ventral scale rows are discontinuous and do not completely crossthe belly region from side to side. In these cases, a right side and a left sidecount is made and given as an average number of rows.

Tail regions:Double caudal whorls: The rings of scales encircling the anterior portion of the tail, that




include the double crest on the dorsal aspect. Double caudal rows are counted,beginning at the rear base of the hind limb, immediately following the last dorsalrow of scales, and ending at the first single tail verticil (crest).

Single caudal verticils: The posterior portion of the tail that is dorsally raised in a singlecrest, following the double caudal whorls. Scale counts are made from the first

single verticil to the tip of the tail, but not including the tip of the tail.Lateral tail region: The sides of the tail, immediately following the hind limbs. Lateral tailinclusions are short series of “extra”

Anal vent: The opening of the cloaca, located on the posterior ventral surfaceimmediately posterior to the hind limbs. The anal vent is surrounded by rings ofsmall scales. The distance from the tip of the snout to the anterior vent generallyequals 50% of the total length of the animal.

Ventral tail or sub-caudal region: The region on the bottom surface of the tailImmediately following the anal vent. Sub-caudal tail inclusions are “extra” scalesthat may occur between the rings of double caudal whorls, on the underside ofthe tail immediately following the hind limbs. These inclusions may be restricted

to the lateral aspects of the underside of the tail, cross the entire underside of thetail, or may be restricted to a midline, extending from the anal vent.

USE OF KEYS

The identification key is composed of paired statements, labeled “a” and “b”,which describe contrasting morphological conditions. These paired statements orcouplets are numbered consecutively.

To identify a live crocodilian, start with couplet 1. Decide whether statement “a”or statement “b”, best describes the specimen in question. After selecting the correctstatement, proceed to the next couplet indicated by the number at the end of thecouplet. Repeat the process until you have completed a couplet ending in a namedspecies. A couplet section may refer you to continue to the couplet immediatelyfollowing, or it may have you skip a large segment of the key and proceed to a coupletfar ahead. Whichever the case is, move only to the couplet indicated. Do not skipcouplets. Individual couplets are not diagnostic alone, but only after the exclusion ofprevious couplets.

All characteristics are externally evident in life, including osteological or skeletalcharacters which may be given. Whenever necessary, the key may refer to a figurewhich should clarify the point in question. Once a specimen has been identified, itshould be further compared with other characteristics listed in the detailed accounts ofthe species for verification, and referral to similar species.




KEY TO FAMILIES OF LIVING CROCODILIANS

1. a) Integumentary sensory organs (ISO’s) present on all body scalation, including the head (Fig. 2. c) -------------------------------------------------------------------------------------- 2

b) ISO’S only present on the head, absent from body scalation ---------Alligatoridae.

2. a) Lower jaw joined (mandibular symphysis) to the level of the fourth, fifth, seventh,or eighth mandibular teeth (Fig. 7) ------------------------------------------Crocodylidae

b) Lower jaw (mandibular symphysis) joined to the level of the 14th. or 21 st to 23rd..mandibular teeth (Fig. 5, 6) -------------------------------------------------------- Gavialidae

KEY TO THE SPECIES Of LIVING CROCODILIANS

1. a) Enlarged fourth mandibular tooth of the lower jaw received into an internal socket

in the upper jaw, hidden when the jaws are closed (Fig. 2a). Ventral scales lackISO’s (Fig. 2c)------------------------------------------------------------------------------------ 2 b) Enlarged fourth mandibular tooth received into an external notch on the

sides of the upper jaw, visible when the jaws are closed (Fig.2a). Ventral bodyscales have ISO’s (Fig.2c) ------------------------------------------------------------------- 7

2. a) Iris of eyes dark brown, red, or orange------------------------------------------------------3 b) Iris of eyes other than dark brown, red, or orange----------------------------------------5

3. a) Fewer than 20 teeth/sockets on each side of lower jaw----------Al l igator sinensis b) More than 20 teeth/sockets on each side of lower-------------------------------------- 4

4. a) Ear coverlets black or dark brown (Fig. 3a), contrasting with lighter color of thecranial table. Two, occasionally three, lumbar rows of dorsal scales above thehind limbs (Fig. 3b)----------------------------------------------- Paleosuchu s trigon atus .

b) Ear coverlets colored similar to the cranial table. Minimum of four longitudinallumbar rows of dorsal scales above hind limbs------ Paleosuchu s palpebrosus .

Fig.2a No externalnotch for tooth.

Internal socket.

Fig.2b Tooth lies exposed inexternal notch(see also Defin ing Terms, p6.)

Fig.2c ISO’s

Tail

Head




Fig. 3a Color of coverlets Fig. 3b Two or four scales

5. a) Body color black overall, without yellow flecks. Body and tail with distinct narrowyellow or white cross-bands, at least on the sides, particularly in the young-------6

b) Body color dark or brown overall, often with numerous yellow flecks; or bodyoverall yellow, or green, with diffused dark cross-bands and flecks, never distinctwhite cross-bands (See Key to Caiman, p. 17)---------------------------------- Caiman .

6. a) Three or more large dark blotches on the sides of the jaws (Fig. 4a). Dorsal neckcolor behind skull, light tan or beige (Fig. 4b). ----------------Melanosuch us niger .

b) Jaws light or dark, but lacking large dark blotches (Fig. 4c). Neck color behindskull black (Fig. 4d).------------------------------------------- Al l igator mississip piensis

7. a) Mandibular symphysis extending caudad to the level of the 21st. or 23

rd.

mandibular teeth (Fig 5) ------------------------------------------ Gavi ali s gangeti cu s

Fig. 4a. M. niger. Beige neckJaw blotches

Fig. 4c A. mississippiensis

Black neck; no jaw blotches

4b 4d

Cranial table




b) Mandibular symphysis not as above-------------------------------------------------------- 8

8. a) Mandibular symphysis extending caudad to the level of the 14th or 15th

mandibular teeth (see Fig. 5) (Fig 6) ----------------------------- Tomistoma schlegel ii

b) Mandibular symphysis not as above-----------------------------------------------------------9

9. a) Mandibular symphysis extends caudad to level of the fourth or fifth mandibularteeth (Fig. 7a) -------------------------------------------------------------------------------------12

b) Mandibular symphysis extends caudad to the level of the sixth, seventh, oreighth mandibular teeth (Fig 7b) (See Fig. 5) -------------------------------------------- 10

Skull.

Lower jaw tooth 21-23

Lower jaw

tooth #1

Fig. 5

Lower jaw tooth

14 - 15

Lower jaw

tooth 1

As seen in life.

Lower jaw tooth 21-23

Fig. 6




10. a) Nuchal scales arranged in longitudinal rows, continuous with the dorsalscales (Fig. 8a) ------------------------------------------------ Crocodylu s cataphractus

b) Nuchal scales arranged in a cluster of four to six scales (Fig. 8b), separatedfrom the dorsal scales by a transverse row of fewer than four, usually small,scales or an expanse of soft skin ----------------------------------------------------------11

11. a) Nuchal scales formed in a cluster, closely followed by one or two rows of fewerthan four, usually small scales, forming a continuous series with the dorsal scales(Fig 9a) (see also Fig. 8a). Ventral scales in 22 to 24 transverse rows --------------

------------------------------------------------------------------------------ Crocody lus johnson i b) Nuchal scales widely separated from the dorsal scales by an expanse of softskin (Fig 9b). Ventral scales in 25 to 27 transverse row ---------------------------------

-------------------------------------------------------------------------Crocodylu s intermedius

1s

Dorsalrow

Pairednuchal

rows

2 34.

65

1

Fourth to fifth tooth

Sixth to seventh tooth

Fig. 7aFig. 7b

Fig. 8a Fig. 8b




+

Fig. 9a. C. johnsto ni Fig. 9b. C. acutu s

12. a) All ventral surfaces mottled or uniformly dark brown or black. Nuchal scales arearranged in longitudinal rows, continuous with the dorsal scales (see Fig. 8a) – ------------------------------------------------------------------------------------------------------ 13

b) Ventral body surfaces uniformly white or cream, with or without lateral dark barsbarely entering the ventral scales or dark sub-caudal tail spots. Nuchal scalesarranged in a cluster of four to six scales (See Fig. 7a) ------------------------------14

13. a) Interior of the mouth (palate) uniformly white or cream, usually two transverserows of post-occipital scales (Fig. 1) ------------ Osteolaemus tetraspis o sborn i

b) Interior of the mouth (palate) darkly mottled anteriorly, yellow or creamposteriorly, usually, a single transverse row of post-occipital scales (Fig.1) ------------------------------------------------------------------- Osteolaemus tetraspis tetraspis

14. a) A distinct pre-orbital ridge, extending well onto the snout from each eye

(Fig. 10a); or irregular scales extending caudad from the anal scales in an

irregular line, through the first several rings of sub-caudal tail whorls(Fig 10b) -----------------------------------------------------------------------------------------15

b) No pre-orbital ridge, or a short ridge extending onto the snout less than a thirdof the distance to the nostril openings. Irregular scales not extending caudadfrom the anal vent through the first several sub-caudal tail whorls -------------- -16

Dorsals

Nuchal cluster




15. a) Ventral scales in 28 or fewer transverse rows ------------------------------------------17b) Ventral scales in 29 or more transverse rows -------------------------------------------18

16. a) Transverse dorsal scale rows irregular. Rows composed of from two to sixscales each, separated by soft skin, with scales often missing within arow (Fig. 11a) -----------------------------------------------------------------------------------19

b) Transverse dorsal rows regular. First row composed of two to four scales,followed by six to 12 rows of six or more scales, again followed by several rows

of four scales (Fig 11b) -------------------------------------------------------------------------- 20

17. a) Nuchal scales separated along the midline by a space of soft skin -------------------------------------------------------------------------------------- Crocodylu s novaeguineae

b) Nuchal scales not separated, scales in medial contact with each other ----------------------------------------------------------------------------------- Crocody lus mindorens is

Fg. 10a Fig. 10b

Fig. 11a Irregular rows, scales “absent.” Fig. 11b Highly uniform rows




18. a) A series of irregular scales extending caudad from the posterior anal vent (as inFig. 9b) and continuing through the first several rings of sub-caudal tail whorls.two pairs of post-occipital scales always present (Fig 1) -Crocodylu s siamensis

b) No series of irregular scales extending caudad from the posterior anal vent.

Post-occipital scales usually absent, or represented by only one or two smallScales -------------------------------------------------------------------Crocody lus po rosus

19. a) Width of the mandible at the posterior margin of the mandibular symphysis(Fig. 12a) equal to or greater than the length from (Fig. 12b) the posterior marginof symphysis to the tip of the mandible ----------------------------------------------------22

b) Width of the mandible at the posterior margin of the mandibular symphysis(Fig. 12a) less than the length (Fig. 12b) from the posterior margin to the tip ofthe mandible -------------------------------------------------------------- Crocody lus acutus

20. a) More than 30 transverse ventral rows from the ventral collar to the anal vent.Toes of the front feet extremely short and stump -----------Crocody lus rhom bi fer

b) Thirty or fewer transverse ventral rows, toes elongate and well developed ----21

21 a) Fewer than 26 transverse ventral rows. Flank scalation uniform in size, round or oval, in longitudinal rows ---------------------------------- Crocodylu s novaeguineae

b) Fewer than 26 transverse ventral rows. Flanks with one or two rows of enlargedkeeled scales. -------------------------------------------------- Crocody lus mindorens is

c) From 26 to 30 transverse ventral rows.. Nuchal cluster not medially divided orseparated; in contact along the midline ------------------------Crocodylu s ni lot icus

22. a)Sub-caudal tail whorls interrupted by irregular scales or groups of scalesbetween the rings (Fig. 13). Young and adults dark greenish-yellow or uniformdark olive-drab -------------------------------------------------------Crocodylu s moreleti i

b) Sub-caudal tail whorls arranged in uniform rings following one upon theother without interruption. Young and adults brown or yellow-brown with darkbrown bars and spots--------------------------------------------------------------------------23

ab

Fig. 12




23. a) Ventral collar scales (Fig. 1) not distinctly enlarged, majority of transverse dorsalrows include four scales ------------------------------Crocodylu s palustr is palustr is

b) Ventral collar scales distinctly enlarged, majority of transverse dorsal rows

include six scales ---------------------------------------Crocodylu s palustr is kimbula

Fig. 13

Dark scales indicate where random scaleinclusions occurr between rows of sub-

caudal scales.

Anal vent




Fig 14

Fig 15

KEY TO THE SPECIES OF THE GENUS Caiman

Key characteristics of the genus Caiman Upper surface of the upper eyelid raised in a pointed elevation (Fig 14).

A transverse bony ridge extending from the anterior point of each orbit, across the snoutbetween the eyes, to the anterior point of the other eye (Fig 15).

Color patterns indicated are typical for juvenile and adult caiman. Recent hatchlings ofmembers of the genus Caiman, less than 25cm in length, tend to be colored brownoverall with darker cross-bands, and have several faint bars on the sides of the upperand lower jaw. Hatchling patterns are lost within weeks after hatching. Caiman yacare is an exception, in that hatchlings are markedly colored with the adult patterns, including




bold jaw markings, at birth. Very old adult caiman tend to become overall unicolor withage, but tend to retain ventral, jaw, flank and lateral tail patterns. Moistening the animalbefore examination helps enhance colors and patterns.

Key to the Species of Caiman

1. a) Sides of the upper and lower jaws with bold dark blotches (Fig 16a, b); marbleddark pattern on the lateral ventral scales (Fig 16c)-------------------Caiman yacare

b) Sides of the jaws unmarked (Fig 17a) or with faint or diffused spots or flecks (Fig 17b); lateral ventral scales un-patterned (Fig 17c) or with short dark bars andspots (Fig 17d) --------------------------------------------------------------------------------------2

Fig. 16

a) Bold

b)

c)

b) DiffusedC. latirostrisC. c. crocodilus

Fig. 17a None Fig. 17b Diffused




Fig. 17c Fig. 17d

Fig. 18

2. a) Snout tapered (trapezoidal) and narrowing from the orbits to the tip of the snout. A transverse bony ridge extends only from the anterior of one orbit to anterior of

the other orbit (see Fig. 15) ----------------------------------------------------------------------- 3

b) Snout wide and blunt (square and box-like), nearly as wide as the distance fromthe orbit to the tip of the snout. A transverse bony ridge extends from the anteriorof one orbit to the other orbit, continuing as an elevation, laterally on the snout tothe sides of the jaws (Fig. 18) ------------------------------------------- Caiman latirost r is




Fig. 19b Fig. 19a

Fig. 20a

3. a) Jaw markings absent. Overall body color nearly uniform dusky brown with obscurefaint or absent cross bands. Faint cross-bands on the back and lateral tail (Fig.

19a). Flank scales in three to four uniform rows of oval, slightly keeled scales,alternating with rows of small scales (Fig. 19b).. Lateral ventral scales un-patterned (Fig. 17c) --------------------------------------------- Caiman crocodi lus fuscu s

b) Several diffused small spots and blotches on the sides of the lower jaw (Fig17b). Flanks scales as in Fig. 19b. Ventrals often with small diffused spots.Overall body color drab yellow or brownish green with diffused, and obscurecross-bands and flecks (see species account)------------------------------------------------------------------------------------------------ Caiman c. chiapasius (= Caiman c. fuscu s) .

c) Without jaw pattern, or with one or two faint diffused shadow like blotches on thelower jaw only (Fig. 17b). Lateral ventral scales un-patterned or with short diffused

marks. Body color yellow, green or yellow-brown with dark cross-bands and flecks(Fig 20a). Flank scales composed of two to four rows of large, often greatlyelongated, heavily keeled scales, surrounded by an expanse of softcreased skin (Fig 20b). -----------------------------------Caiman crocodi lus crocodi lu s

2

3




Fig. 20b

Elongated keeled scales

Dorsal scales

1Enlarged row2 Enlarged row3 Enlarged row

Ventral scales

Skin

Mid-body




SPECIES ACCOUNTS FOR LIVING CROCODILIANS

Order: Crocodylia

Family: Alligatoridae

Al l igator mississippiensis (Daudin)American alligatorCONTENT: No subspecies are recognized.

SIZE: Maximum size suggested up to 570cm. Average size 200cm to 300cm.Hatchlings 22cm to 27cm

COLORATION: Black above with yellow or white cross-bands on the back and tail. Lightcolored variations on the sides of the body and tail, extending down the flanks to formbars. Light flecks usually on the sides of the head, and neck. Sides of the jaws light

colored. Dark bars on the lateral ventral scales. Belly often mottled with gray or black.Juveniles colored as the adult. Old animals usually uniform gray-black.

CONFORMATION: Head and snout rugose. Snout wide, flat and blunt; length 1 to 1.5times the width at the base. Integumentary sense organs (ISO’s) found on the headonly.

SCALATION:Post-occipitals: 2 to 3 transverse rows of irregularly arranged small keeled scales.Nuchals: 3 to 4 transverse rows of 2 to 4 scales, continuous with the dorsal scalesDorsals: 17 to 18 transverse rows of 8 to 10 scales.Double crest caudal whorls: 17 to 19.Flanks: Scales on the sides of the body round to oval, nearly uniform in size, each nextto the other in loose longitudinal rows.Single crest caudal whorls: 18 to 21.Ventral collar: a single transverse row of enlarged scales.Ventral scales: 30 to 35 transverse rows.Lateral tail: Inclusions of scales often occur between the anterior tail whorlsSub-caudal scale rows: uniform uninterrupted rings.

Anterior lateral tail: Typical have two to six or more inclusions.Flank scalation: a continuous pattern of like size, un-keeled, round and oval scales(Fig. 21).




Fig. 21

DENTITION: Fourth mandibular tooth received into an internal socket in the upper jaw.Mandibular symphysis extends to the level of the fourth or fifth mandibular teeth. Theteeth are short and thick. Posterior teeth rooted into a common elongated socket.Formula: 5 + 13-15.

19-20

SIMILAR SPECIES: Melanosuchus niger : has from three to five large dark spots on thesides of the jaws. Neck at base of head tan or beige. Alligator sinensis: fingers are notwebbed; gray or gray-green above with faint gray and yellow cross-bands. Old adultsuniform dark gray. Fewer than 28 transverse ventral rows.

DISTRIBUTION: From Corpus Christi, Texas, east to Florida; north to Tyrrell County,North Carolina; the Mississippi River system, Arkansas and formerly southeastOklahoma.

COMMENTS: Spelling of the species taxon corrected to “mississippiensis” from original“mississipiensis”.

REFERENCES: Boulenger, 1889; Cope; Daudin, 1802; Kellogg, 1929; Mook, 1921b;Oliver, 1958; Schmidt, 1922,1944; Thorbjarnarson, 1992; Wermuth, 1953; Wermuth andMertens, 1961; Werner, 1933.




Al l igator sinensis FauvelChinese alligatorCONTENT: No subspecies are recognized.

SIZE: One of the dwarf species of crocodilians. Males are on the average smaller than

females. Average adult size is 100 cm to 130 cm; maximum size, approximately 200cm. Hatchlings, 15cm to 25cm.

COLORATION: Young adults are dark gray over all with indistinct lighter cross-bandson the back, sides and tail. Ventrals are mottled with dark and light gray. The head islightly speckled with yellow or white. Some retain a gray-green ground color. Hatchlingsare dark brown or black with white or orange markings; a brightly colored “X” crossesthe snout immediately in front of the eyes. There are gray specks on the sides of thelower jaw. The hatchling patterns are lost by the end of the first year and replaced withvivid pre-adult yellow-green and gray colors with yellow or white cross-bands. Adultsand old adults become uniform dark gray, nearly black. The eyes are large, dark brown,

and appear bovine-like. Flanks: a continuous pattern of like size, un-keeled, round andoval scales.

CONFORMATION: The head is massive and wide. The snout is round, wide, and blunt,slightly longer than wide at the base. The orbits are not connected anteriorly by atransverse bony ridge. Two or more poorly developed ridges may extend from theanterior orbits well onto the snout, and usually become overgrown with age, producingan almost smooth surface. The upper eyelids are rugose with light colored welldeveloped upper and lower striated “eyelashes”. The fingers of the front feet are notwebbed. ISO’s are absent on the body.

SCALATION:Post-occipitals: a single transverse row of 6 to 8 separated keeled oval scales,separated from the nuchals by 2 isolated oval scales.Nuchals: 3 pairs of enlarged scales arranged in 2 longitudinal rows, in contact with eachother, and closely followed by the first transverse dorsal row of scales.Dorsals: 16 to 17 transverse rows of 6 scales.Double crest caudal whorls: 16 to 17.Flanks: 5 to 7 longitudinal rows of raised, equal sized, round scales. Smaller, randomlyplaced, bead-like scales often occur in the space between the rows.Single crest caudal whorls: 16 to 17.Ventral Collar: a single enlarged transverse row of scales.Ventral scales: 25 to 27 transverse rows.Lateral tail: unremarkableSub-caudal scale rows: uniform uninterrupted rings,

Anterior lateral tail: Usually none.Flank scalation: a continuous pattern of like size, un-keeled, round and oval scales(Fig. 22).




Fig. 22.

DENTITION: Fourth mandibular tooth received into an internal socket in the upper jaw.Mandibular symphysis extends to the level of the fourth or fifth teeth. The teeth are thickand short. Those on the rear portions of the mandible and maxillary are rooted in a

common elongated socket. Formula: 5 + 13-14.18-19

SIMILAR SPECIES: Alligator mississippiensis: fingers of the front feet are extensivelywebbed, ventral scales in more than 29 rows. Melanosuchus niger : median ventralsurface immaculate white. Paleosuchus: red-brown head and body with extensivebrown blotching on the ventral surfaces; only four teeth on each side of thepre-maxillary. Osteolaemus: fourth mandibular teeth are exposed in an external notch;ISO’s are present on the head and body scalation.

DISTRIBUTION: Historically Inhabited the alluvial plains and lowlands along rivers andstreams, principally the basin of the Chang Jiang (Yangzi River), extending northward to

a portion of the eastern Huang He (Yellow River). However, the distribution of thisspecies, one of the most endangered crocodilians, has been greatly reduced to mostlycaptive populations with few remaining in altered habitats

COMMENTS: Little taxonomic confusion exists concerning this species. However, muchof the life history data is confounded in folklore and superstition

REFERENCES: Barbour, 1922; Boulenger, 1889; Chu Cheng-kuan, 1957; Fauvel,1879; Hsiao, 1934; Mook, 1923; Thorbjarnarson, 1992; Wermuth, 1953; Wermuth andMertens, 1961.




Caiman crocod i lus (Linnaeus) CaimanCONTENTS: Three races of Caiman crocodilus are commonly recognized: Caiman c.apaporiensis, Caiman crocodilus crocodilus, Caiman c. fuscus.

CONFORMATION: All forms. Integumentary sense organs (ISO’s) found on the headonly. The fingers are not webbed. A well developed transverse bony ridge connects theorbits, anteriorly. The upper eyelids are raised into a high point or tubercle (Fig. 14, 15).

COMMENT: Five races comprising the genus Caiman have historically beenrecognized. Caiman crocodilus crocodilus, Caiman c. apaporiensis, Caiman c. fuscus,Caiman c. yacare, and Caiman latirostris. Caiman sclerops is considered a synonymfor Caiman crocodilus Caiman c. chiapasius (Bocourt, 1876) (= C. c. fuscus) describedfrom the Choco of Colombia, Gorgona Island, Central America, and Mexico is notconsidered here. A review of the Caiman taxonomic literature is presented in King andBurke (1989) The taxon Caiman yacare (= Caiman c. yacare) is included here as a

separate species. In recent years, Caiman have been the subject of extensive fieldstudies, molecular systematics, distribution, and morphological studies.

The Caiman complex requires special comment, in that a number of additional sub-species level taxa have been cited in literature over the past 30 years (see individualspecies accounts), based, for the most part on commercial skins of undocumentedorigins and conflicting distributional limits. These taxa include: Caiman c.matogrossiensis and C. c. paraguayensis (Wermuth and Mertens, 1977). They arecontinued, however, in the international CITES Identification Manual (1981, 1983), forthe identification of crocodilians and their derivatives.

Recent field studies and subsequent DNA genetic analysis of blood and tissue samples,and statistical analysis of morphological data, combined with zoogeographical analysisof geological and hydrological parameters separating the species populations, did notconfirm the existence of these taxa (Brazaitis et al., 1997a, b). These studies suggestthat the genus Caiman is more properly composed of the taxa. Caiman crocodilus,Caiman fuscus (which may include Caiman c. chiapasius), Caiman latirostris (= Caimanl. chacoensis) , and Caiman yacare, each comprising a single distinct andzoogeographically separated species.

DNA comparison of tissue samples of Caiman c. apaporiensis type specimens collectedby Dr. Federico Medium in the upper drainages of the Rio Solimões in Colombia, withCaiman crocodilus crocodilus samples from throughout Amazonia in Brazil, alsosuggest they are one and the same species. Caiman crocodilus crocodilus.

REFERENCES: Amato,1993; Amato et al., 1998; Bocourt, 1876; Boulenger, 1889;

Brazaitis, 1986; Brazaitis et al., 1993 (unpublished report); Brazaitis et al., 1997a,

(unpublished report); Brazaitis et al.,1997b; Brazaitis et al.,1998

a,b; De Carvalho, 1955;

CITES Identification Manual, 1981; Densmore and Owen, 1989; Densmore and White,1991; Frair and Behler, 1983; Fuchs, 1971, 1974; Fuchs et al., 1990; Hagmann, 1906;




King and Burke,1989; King and Roca, 1989 (unpublished report); Linneaus, 1758;Medem, 1955, 1960, 1981, 1983; Medem and Marx, 1955; Mertens, 1943; Mook,

1921b; Muller and Hellmich, 1936; Schmidt, 1928

a; Schmidt, 1928

b; Scott et al., 1988,

1990; Spix, 1825: Thorbjarnarson, 1992; Wermuth, 1953, 1986; Wermuth and Fuchs,1978; Wermuth and Mertens, 1961, 1977; Werner, 1933.

Caiman crocodi lus crocodi lu s (Linnaeus) Common caiman

SIZE. Maximum size, 250 cm; Average adult, 150 cm to 200 cm; hatchlings, 20 cm to25 cm.

COLORATION. Dorsally olive green with numerous dark brown or black flecks on thehead, body, and tail. Dark brown or black cross-bands on the back and tail. Ventralsurfaces are un-patterned, uniform cream, or yellow. No large dark blotches on the

sides of the jaws. Hatchlings display a series of faded bars on the sides of the jaws, nolonger visible at about 35 cm total length.

CONFORMATION. Snout moderate and slightly elongate, longer than its width at thebase.

SCALATION.Post-occipitals: 2 to 3 relatively regular transverse rows of 6 to 8 enlarged scales.Nuchals: 4 to 5 transverse rows of scales. Each row closely followed by the next. Twoor more rows containing 4 scales each. Nuchal scales are continuous with the dorsalrows.

Dorsals: 18 to 19 transverse rows of 8 to 10 scales.Double crest caudal whorls: 12 to 13.Single crest caudal whorls: 20 to 22Ventral collar: a single enlarged transverse series of scales.Ventral scales: 20 to 24 transverse rows.Sub-caudal scale rows: Uninterrupted uniform rings.

Anterior lateral tail: Typical without anterior lateral tail inclusions, rarely one to twoinclusions.Flank scalation: Typical are two to three rows of enlarged, heavily keeled, elongatedscales between the dorsal and ventral scalation. The upper row is composed of thelargest scales. Typically, all rows are elongated. However, lower second or third scale

rows of Venezuelan populations tend to retain distinct keeling but may be round toelongate. All rows are widely separated from each other by creased soft skin andrandom small scales (Fig. 23).




Fig. 23

DENTITION: As in Caiman c. crocodilus

SIMILAR SPECIES: Caiman yacare bear large dark blotches on the sides of the jaws,Caiman latirostris: both. Caiman c. fuscus: dorsally yellow-brown or brown overall,without dark brown or black dorsal speckling; no dark cross-bands on the back in

specimens 90 cm or more in length; double crest caudal whorls 13 to 14, snout mod-erate. Caiman c. apaporiensis: snout elongate and narrow, and anteriorly parallel-sided;bright yellow-brown dorsally.

DISTRIBUTION: Guiana, Venezuela, Trinidad, the Amazon drainage, eastern Colombiato Peru.

COMMENTS. See Caiman.

REFERENCES: See Caiman. Bocourt, 1876; Boulenger, 1889; Brazaitis et al.,1997a,b

;

Brazaitis et al.,1998

a,b

; DeCarvalho, 1955; Hagmann, 1906; Linneaus, 1758; Mook,1921

b; Schmidt, 1928

a; Thorbjarnarson, 1992; Wermuth, 1953; Wermuth and Mertens,

1961; Werner, 1933.

Caiman crocod i lus apapor iensis MedemRio Apaporis caiman

SIZE: Nineteen individuals collected by F. Medem ranged in length from 62 cm to 209cm.

COLORATION: This is a predominantly yellow animal speckled with black or brown.Bright yellow-brown above with black spots and irregular striations, denser on the head.From four to six dark bars on the tail. Extremities dark gray or black.Ventrals: yellow. Some individuals bear yellow scales on the sides of the neck. Youngindividuals reported to be brighter yellow than any other species of caiman.

CONFORMATION: Skull and snout elongate. Snout strikingly elongated, narrow, and




anteriorly parallel-sided, longer than its width at the base (Fig. 24). ISO’s are absent onthe body.

Fig. 24 Caiman c. apapo riensis , FMNH 69828, paratype

SCALATION.Post-occipitals: 2 transverse rows of 6 to 7 scales each.Nuchals: 4 transverse rows of 2 to 4 scales each. The first 2 rows on the type specimenbear 4 scales each. Nuchals continuous with the dorsal scale rows.Dorsals: 19 transverse rows, the widest containing 9 scales.Double crest caudal whorls: 13.Single crest caudal whorls: not reported.Ventral collar-,a single transverse series of enlarged scales.Ventral scales: 20 to 24 transverse rows.

Sub-caudal scale rows: not reported. Anterior lateral tail: Unreported.Flank scalation: Appears to be similar to Caiman c. crocodilus (Fig. 23 ).

DENTITION. As in Caiman crocodilus crocodilus.

SIMILAR SPECIES. Caiman c. crocodilus. dark olive-green above, with dark brown orblack flecks; snout slightly elongate, not anteriorly parallel sided. Caiman c. fuscus:yellow-brown, brown, dusky brown above; no dark flecks or spots on the dorsal surfacesof the body; extremities yellow, yellow-brown as is the body. Caiman yacare: large darkblotches on the sides of the jaws and snout. Caiman latirostris: yellow body coloration,

from three to five large diffused blotches on the sides of the jaws in some individualsfrom the Choco of Argentina.

REMARKS: Little is known of this crocodilian's life history or ecological relationships.Isolated by waterfalls and rapids.

DISTRIBUTION: Reported to be found only in the upper Rio Apaporis, between the.falls of Jirijirimo and Puerto Yiviya, a distance of about 200 kilometers in Colombia.




COMMENTS: It is important to note that preliminary molecular and morphologicalstudies suggest that this species may be a synonym for Caiman c. crocodilus (Brazaitiset al., 1997). However, Fuchs et al. 1990, report skins of this species in commercialtrade. Type and paratype specimens are deposited in the Field Museum of Natural

History. Additional specimens in the Instituto de Ciencias Naturales Bogota, Colombia.

REFERENCES: See Caiman. Brazaitis et al.,1997a,b

; Fuchs et al., 1990; Medem, 1955,

1960, 1981, 1983; Wermuth,1986; Wermuth and Mertens, 1961.

Caiman crocodi lus fuscu s (Cope) Dusky or brown caiman

SIZE. Maximum size approximately 225cm. Average adults 125cm to 175cm.Hatchlings, 21cm to 26cm.

COLORATION: Adults light brown, olive brown or light yellow above. Dark cross-bandsvisible on the sides of the tail only. Ventrals, uniform cream or yellow. Old adults losealmost all signs of cross bands and appear uniformly dark dusky brown or olive brown.Hatchlings, light brown or yellow with dark brown cross-bands on the back and tail. Fiveto seven faint dark bars usually evident on the sides of the jaws, no longer visible onanimals 35 cm or larger.

CONFORMATION: Snout triangular, gradually converging sides. Snout equal to orshorter than its width at the base. ISO’s are absent on the body.

SCALATION.Post-occipitals: 3 to 4 transverse rows of 6 to 10 slightly enlarged irregularly arrangedscales.Nuchals: 4 to 5 transverse rows of 2 to 4 scales, continuous with the dorsal scales.Dorsals: 17 to 18 transverse rows of 8 to 10 scales.Double crest caudal whorls: 13 to 14.Single crest caudal whorls: 23 to 24.Ventral collar: a single transverse row of enlarged scales,Ventral scales: 22 to 24 transverse rows of 12 to 15 scales each.Sub-caudal scale rows: uniform uninterrupted rings.

Anterior lateral tail: Typical without anterior lateral tail inclusions, rarely one inclusions.

Flank scalation: Typically three to four rows, rarely five rows, of oval poorly keeled,enlarged scales. The upper row is composed of the largest scales. Flank scales arearranged in uniform parallel rows. Each row is separated from the other by a uniformrow of small bead-like scales (Fig. 25)




Fig. 25

DENTITION. See Caiman c. crocodilus.

SIMILAR SPECIES: Caiman c. crocodilus: dark olive-green above, flecked with dark

brown or black on the head, body and tail; dark cross-bands visible on the back and tail;double caudal whorls: 12 to 13; snout, slightly elongate, slightly longer than its width atthe base. Caiman yacare, Caiman latirostris: from four to five large blotches on thesides of the jaws. Caiman c. apaporiensis: snout strikingly narrow and elongate,anteriorly parallel-sided; dark cross-bands evident on the back and tail; double caudalwhorls 13; some individuals bear orange scales on the sides of the neck; dorsal color asCaiman c. fuscus; numerous dark spots on the back, sides, and head.

DISTRIBUTION: Southern Mexico, Panama through Central America, Colombia andthe river systems Magdalena, Sinu, and Rio Atrato (Figure 10).

COMMENTS: Caiman c. chiapasius is sometimes used to identify members of this racefrom the Choco of Colombia, Gorgona Island, Central America, and Mexico. That taxonis not generally accepted as valid.

REFERENCES.: See Caiman. Bocourt, 1876; Brazaitis et al.,1997a,b

; Brazaitis et

al.,1998a,b

; Cope, 1868; Medem, 1955, 1960,1962, 1968, 1969, 1981, 1983; Miguel

Alvarez Del Toro, 1967, in lit.; Schmidt, 1928a; Thorbjarnarson, 1992; Wermuth, 1953;

Wermuth and Mertens, 1961; Werner, 1933.

Caiman yacare (Daudin) Yacare caimanCONTEN: No subspecies are recognized.

SIZE:. Adults 250cm are common. Hatchlings average 20cm to 30cm.

COLORATION: This is a predominantly dark animal, speckled with yellow. Adultsusually appear black above. The speckled young become progressively darker with age




and assume the adult coloration at about 100cm in length. Young and adults arecharacterized by three to five large dark blotches on the sides of the jaws. Ventral whiteor slightly cream, patterned along the lateral margins with dark marbling, extending ontothe ventral scales from the sides. Some individuals bear reddish or orange marbling onthe sides of the body or neck. Such individuals probably led K. P. Schmidt to inquire into

the identity of the "red jacare" of Paraguayan report.

CONFORMATION: Snout and skull triangular, without parallel sides. Snout slightlylonger than the width at the base. The first two teeth of the mandible as well as thefourth mandibular teeth invariably pierce the snout of adult animals. ISO’s are absent onthe body. The fingers lack webbing.

SCALATION.Post-occipitals: 3 to 4 irregular transverse rows of enlarged scales.Nuchals: 4 to 5 transverse rows of scales. The first 2 to 3 rows composed of 4 scaleseach, latter rows of 2 to 3 scales each, continuous with the dorsal scales.

Dorsals: 17 to 18 transverse rows of 8 to 10 scales each.Double crest caudal whorls: 13 to 15. Single crest caudal whorls: 17 to 19.Ventral collar: a. single transverse series of enlarged scales.Ventral scales: 22 to 25 transverse rows of 14 to 16 scales each.Sub-caudal scale rows: uniform uninterrupted rings.

Anterior lateral tail: Frequently two to six scale inclusions.Flank scalation: Typical are four to six rows of round or slightly oval enlarged scalesThe uppermost row is composed of slightly larger scales, and is most heavily keeled.Remaining rows are composed of poorly keeled, or smooth un-keeled round or slightlyoval scales. Flank scale rows are closely associated to each other, separated mostoften by a narrow chain like pattern of soft creased skin (Fig. 26).

Fig. 26

DENTITION: See Caiman c. crocodilus.

SIMILAR SPECIES: Caiman latirostris: yellow, ventral collar composed of twotransverse rows of slightly enlarged scales; snout length equal to or one and one-fourth




times the width at the base; only the first transverse row of nuchal scales composed offour scales. Melanosuchus niger : upper eyelid flat above, or only a feeble raisedprojection or tubercle; median two longitudinal rows of dorsal scales elevated along thevertebral line.

DISTRIBUTION: Southern Brazil, Paraguay, Bolivia. The river systems of Parana,Paranaiba, north to the Sao Francisco; the whole of the Mato Grosso; the rivers ofItenez and Mamore .

COMMENTS: See Caiman. Caiman c. matogrossiensis and C. c. paraguayensis (= C.yacare) are described from the regions of Brazil, Paraguay and Bolivia, based onundocumented commercial tannery skins.. However these taxa appear to be withoutmerit.

REFERENCES: See Caiman. Amato,1993; Amato et al., 1998; Brazaitis, 1986;

Brazaitis et al., 1993 (unpublished report); Brazaitis et al., 1997a, (unpublished report);

Brazaitis et al., 1997 b; Brazaitis et al.,1998a,b; CITES Identification Manual, 1981;Daudin, 1802; DeCarvalho, 1955; Frair and Behler, 1983; Fuchs, 1971, 1974; Fuchs etal.,1990, Medem, 1955, 1960, 1981, 1983; King and Roca, 1989 (unpublished report);

Medem and Marx, 1955; Mertens, 1943; Mook, 1921b; Schmidt, 1928a; Wermuth, 1953,

1986; Wermuth and Fuchs, 1978; Wermuth and Mertens, 1961, 1977; Scott et al., 1988,1990; Werner, 1933.

Caiman latirostr is (Daudin) Broad snouted caimanCONTENT: Two races of Caiman latirostris are described but not generally recognized:Caiman latirostris latirostris and Caiman latirostris chacoensis.

SIZE: Maximum size probably to 300cm. Adults up to 250cm have been recorded.Hatchlings average 22cm to 30cm.

COLORATION.: Adults yellow, yellow brown or olive green above, densely flecked withdark brown or black. Wide dark cross-bands on the tail, sometimes slightly evident, onthe back. Old adults appear uniform yellow brown with faint bars visible on the sides ofthe tail. Hatchlings and juveniles, yellow brown above with dark brown or black flecksand cross-bands on the back and tail, flecks on the head. All stages are uniform creamor yellow on the ventral surfaces with faint marbling on the lateral ventral scales. Fromthree to five faint blotches on the sides of the jaws.

CONFORMATION: Skull and snout low, wide, and extremely blunt.. Snout length equalto or wider than length to tip of snout. The raised transverse bony ridge which connectsthe anterior points of the orbits is developed into a high inverted "U" shape; extendingfrom the ninth maxillary tooth at the lateral border of the snout, to and across theanterior points of the orbits, then down again to the opposite side of the snout to the




lateral border at the ninth maxillary tooth. The upper eyelids are raised into a point ortubercle, although not as well developed as in Caiman crocodilus. ISO’s are absent onthe body. The fingers are not webbed.

SCALATION.

Post-occipitals: 2 transverse rows of 6 to 8 distinctly enlarged keeled scales.Nuchals: 3 to 4 transverse rows of scales. Only the first of which is composed of 4scales. Each transverse row of scales separated from the next by a short space of softskin. Nuchals continuous with the dorsals.Dorsals: 17 to 18 transverse rows of 6 to 8 scales each, rarely 10.Double crest caudal whorls: 13 to 16.Single crest caudal whorls: 22 to 23.Ventral collar: 2 transverse rows of slightly enlarged scales.Ventral scales: 26 to 30 transverse rows of 12 scales each, average 27 to 28 rows.Sub-caudal scale rows: uniform uninterrupted rings.

Anterior lateral tail: Typical have two to five scale inclusions.

Flank scalation: Typically three to four rows, rarely two rows, of extremely large,elongated, and heavily keeled scales. The upper row is composed of very largeprominent scales, arranged in a row. Rows are widely separated from the dorsal scalesand from each other (Fig. 27).

Fig. 27

DENTITION: Fourth mandibular teeth received into an internal socket in the upper jaw.Mandibular symphysis extends to the level of the fourth or fifth teeth. The posteriormaxillary and mandibular teeth are slightly separated, rooted in nearly unconnected

individual bony sockets. None of the mandibular teeth pierce the upper jaw. Formula: 5+ 12-14, usually 18.17-20 18

SIMILAR SPECIES: Melanosuchus niger : black with yellow or white cross-bands, postoccipitals in three to five transverse rows of small scales; two median longitudinal rowsof scales elevated along the vertebral line; upper eyelids flat, without points or tubercles.Caiman yacare: first two transverse rows of nuchal scales contain four scales each;




each row closely followed by the next. Caiman c. crocodilus, Caiman c. apaporiensis,Caiman c. fuscus: lack large dark blotches on the sides of the jaws. Caiman latirostrischacoensis bears indistinct faded dark blotches on the sides of the jaws. This slightlysmaller race is reported to inhabit the Chaco provinces of Argentina. The ventral scalesaverage 22 to 25 transverse rows.

COMMENTS: In southern Brazil and Paraguay, and possibly southern Bolivia, thisspecies overlaps distribution with Caiman yacare. C. latirostris does not appear tooverlap M. niger distributions in Brazil or Bolivia, as it occurs in more temperate orhigher coastal habitats. Paleosuchus palpebrosus occurs in the same habitats in someregions. .

REFERENCES: See Caiman. Boulenger, 1889; Brazaitis et al.,1997a,b

Brazaitis et

al.,1998a,b

; Cohen and Gans, 1970; Daudin, 1802; DeCarvalho, 1955; Freiberg and

Carvalho, 1965; Mook, 1921 a,b

; Muller and Hellmich, 1936; Schmidt, 1928a; Wermuth,

1953; Wermuth and Mertens, 1961; Werner, 1933; Yamashita et al., 1992.

Melanosuch us niger (Spix) Black caiman,CONTENT. No subspecies are recognized.

SIZE: One of the largest members of the Alligatoridae. Males may reach 500cm inlength. Average adult 300cm, to 400cm, hatchlings about 30cm.

COLORATION: One of the few species which retains most of the juvenile colorationthroughout life. Jet black above with narrow yellow cross-bands on the body and tail.Neck region, immediately behind the skull, is light tan or yellow. From three to five largedark blotches on the sides of the lower and upper jaws. Ear coverlets black, as is themedian dorsal surface of the snout. Median ventral surfaces immaculate white,bordered by jet black. There appears to be some dichotomy among populations. Whitecoloration restricted to the mid-ventral portions of the abdominal region in specimensfrom Brazil; all white with black markings barely extending onto the lateral ventral scalesin those from the south and Bolivia.

CONFORMATION: The head is massive, wide, and high. Snout is slightly pointed “boatshaped”, slightly longer than the width at the base. Orbits are large, connectedanteriorly by a feeble transverse bony ridge, with a deep concave depression in front ofeach orbit. Four oblique elevations extend to the lateral margins of the snout from thepre-orbital region. Upper eyelids are flat, striated, and slightly ossified. The twolongitudinal vertebral rows of scales are elevated to form a visible elevated backboneextending down the back along the mid-line.. ISO’s are absent on the body. The fingersare not webbed.




SCALATION:Post-occipitals: 3 to 5 transverse rows of 6 to 10 slightly enlarged keeled scales.Nuchals: 4 to 5 transverse rows of 2 to 4 scales, continuous with the dorsal scales.Dorsals: 18 to 19 transverse rows of scales.Double crest caudal whorls: 16 to 18.

Single crest caudal whorls: 21 to 24.Ventral collar: 2 transverse rows of slightly enlarged scales.Ventral scales: 25 to 29 transverse rows of from 12 to 14 scales.Lateral tail: Inclusions of 6-8 series of scales frequently occur between the anterior tailwhorlsSub-caudal scale rows: uniform uninterrupted rings.

Anterior lateral tail: Typical have 6-10 scale inclusions.Flanks: 6-8 highly uniform longitudinal rows of round to oval keeled scales, alternatingwith rows of poorly keeled small round to oval scales (Fig. 28).

Fig 28

DENTITION: Fourth mandibular tooth received into an internal socket in the upper jaw.

Mandibular symphysis extends to the level of the fourth or fifth. Teeth thick and short,posteriorly rooted in a common elongated socket. Formula: 5 + 13 -14.

18-19

SIMILAR SPECIES: Alligator mississippiensis: has fingers extensively webbed, nolongitudinal vertebral elevation. Mid-ventral belly region not immaculate white. Caimanyacare: White not restricted to the mid-ventral region. Upper eyelid raised in a point.Ventral scales in less than 26 transverse rows. Caiman latirostris: body color yellow toyellow brown. Mid-ventral belly region not immaculate white. Upper eyelid raised in apoint, a single transverse bony ridge connecting the anterior points of the orbits andextending laterally to the edge of the snout at the ninth maxillary teeth.

DISTRIBUTION: Wild populations continue to be fractionated. Significant populationsexist in localized regions. Although once common throughout the Amazon basin, thisspecies is no longer found in historic numbers anywhere, and has been extirpated frommany previously occupied habitats. Primarily the Brazilian Amazon river basin andtributaries, French Guiana, Guyana, eastern Bolivia, eastern Colombia, Ecuador, Peru.




COMMENTS: This species prefers the quiet waters of oxbow ponds, lagoons, lakes,and open rivers with extensive aquatic vegetation and deep muddy bottoms. Theenvironment is “soft”. Thus, in captivity, individuals easily abrade the feet and chestareas, leading to infection and death. Recovery for this species is difficult.Reproduction takes place at more than 10 years of age, and once populations are

decimated, may not be capable of reestablishing themselves in habitats invaded bymore successful Caiman species.

REFERENCES:. Boulenger, 1889; Brazaitis et al.,1998a,b

; DeCarvalho, 1955;

Hagmann, 1906; Medem, 1963, 1968, in lit; Mook, 1921 a,b

; Schmidt, 1928a, 1944;

Spix, 1825; Thorbjarnarson, 1992; Wermuth, 1953; Wermuth and Mertens, 1961;Werner, 1933.

Paleosuchu s palpebrosus (Cuvier)Dwarf caiman

CONTENT: No subspecies are recognized.

SIZE: Perhaps the smallest of crocodilians. Record is 172cm. Average adult90cm-120cm, hatchlings 15cm-20cm.

COLORATION.: Reddish brown above, ventral surfaces blotched with brown. Someindividuals may appear nearly black, but are usually very dark red-brown on closeexamination. Light areas of the head and body may be light red-brown or yellow. Theear coverlets match the red-brown color of the cranial table and snout. Some large redblotches on the sides of the jaws. The iris of the eyes are usually reddish brown ororange, white surrounding sclera is noticeably evident.

CONFORMATION: The head and snout is square and high, box-like. Snout length oneand one half times the width at the base; unornamented and smooth. The orbits are notanteriorly connected by a transverse bony ridge. The upper eyelids are flat, smooth, andentirely ossified. The post-occipitals and nuchal scales are particularly raised into asharp keel or ridge. All dorsal scales are well-keeled. Transverse rows of ventral scalesoverlap each following row and form an effective bony shield. Lateral longitudinal rowsof ventral scales are keeled. ISO’s are absent on the body. The fingers of the front feetare not webbed.

SCUTELLA TION.

Post-occipitals: 2 transverse rows of small sharply keeled scales. Individuals with onlyone row are not common.Nuchals: 4 to 5 transverse rows of sharply keeled scales. The first row contains 2scales, the second and third rows 3 to 4 scales, followed by I or 2 rows of 2 scaleseach, continuous with the dorsal scales.Dorsals: 18 to 19 transverse rows of 8 scales, reduced to 4 scales over the hind limbs.Double crest caudal whorls: 10 to 11.Single crest caudal whorls: 17 to 21.




Ventral collar: a single enlarged transverse row of scales.Ventral scales: 19 to 24 transverse rows of 16 scales.Sub-caudal scale rows: uniform uninterrupted rings.

Anterior lateral tail: Typical no inclusions.Flanks: Very narrow with usually two rows of heavily keeled elongated scales (Fig. 29 )

Fig. 29

DENTITION: Fourth mandibular teeth received into an internal socket in the upper jaw.Mandibular symphysis extends to the fourth or fifth tooth. The teeth are often tipped withorange, posteriorly rooted in a common elongated socket. Formula: 4+ 14-15.

21-22

SIMILAR SPECIES: Genus Osteolaemus: fourth tooth exposed in a notch. Fourteen to15 teeth on each side of the lower jaw. Genus Melanosuchus: Elevated vertebralcolumn. Five teeth on each side of the pre-maxillary and 17 to 18 teeth on each side ofthe lower jaw. Genus Alligator : has five teeth on each side of the pre-maxillary, more

than 25 transverse ventral rows. Paleosuchus trigonatus: snout elongate and slender,usually only one transverse row of post-occipitals; dorsals reduced to two scales overthe hind limbs; ear coverlets in contrast (dark) to the cranial table, dorsal surface of thesnout -with a median dark stripe.

DISTRIBUTION: The whole of tropical South America, from the Guyana’s in the northand Mexiana Island; south to Sao Paulo, Brazil; the Rio Pastaza in Ecuador to the eastcoast. Probably the most wide ranging species in South America.

COMMENTS: Any confusion concerning this species lies in its ecological relationship toPaleosuchus trigonatus, which together with P. trigonatus make up the genus

Paleosuchus. While P. palpebrosus ranges widely throughout most open river habitatsoutside of deep rainforest, P. trigonatus appears to be mostly restricted to “black water”streams within deep rainforest. Many individuals display overlapping physicalcharacteristics, although P. palpebrosus shows a greater degree of dermal ossification.Interbreeding of the two species under captive conditions, if possible, would be mostinteresting.




REFERENCES: Boulenger, 1889; Cuvier, 1807; DeCarvalho, 1955; Medem, 1958b,

1967; Mook, 1921b; Schmidt, 1928

a; Wermuth, 1953; Wermuth and Mertens, 1961;

Werner, 1933.

Paleosuchu s tr igonatus (Schneider)Smooth-fronted caiman, Schneider’s smooth-fronted caimanCONTENT: No subspecies are recognized.

SIZE: Record is 225cm, male; average adult, 100cm to 130cm; hatchlings, 15cm to20cm.

COLORATION: Dark brown above. Dark brown or black cross-bands on the back andtail. Some individuals laterally tinged with yellow. Cranial table is dark brown, earcoverlets contrasting black or darker brown. A dark brown or black median dorsal stripeon the snout. Ventral surfaces densely blotched with brown. Iris of the eyes reddish

brown or orange, white surrounding sclera noticeably evident.

CONFORMATION: Cranial table square and high. Snout elongate, pointed, length 1.5times the width at the base. Orbits not anteriorly connected by a transverse bony ridge.Upper eyelids flat, entirely bony. Post-occipital and, nuchal scales sharply keeled,raised. Lateral longitudinal rows of dorsal scales keeled, median rows flattened, feeblykeeled. Lateral longitudinal rows of ventral scales keeled. Transverse rows of ventralscales overlap the following row, forms an ef fective bony shield. ISO’s are absent on thebody. Fingers are not webbed.

SCALATION.

Post-occipitals: usually a single transverse row of sharply keeled enlarged scales. Asecond row of smaller enlarged keeled scales not uncommon. Nuchals: 4 to 5transverse rows of 2 scales, continuous with the dorsal scales.Dorsals: 17 to 18 transverse rows of 6 scales, reducing to 2 to 3 scales over the hindlimbs.Double crest caudal whorls: 9 to 10. Single crest caudal whorls: 17 to 19.Ventral collar: a single enlarged transverse row of scales.Ventral scales: 18 to 24 transverse rows of 10 to 12 scales.Sub-caudal scale rows: uniform uninterrupted rings.

Anterior lateral tail: Typical no inclusions.Flanks: Fairly wide flank regions with coarse grainy skin and randomly spaced enlarged

heavily keeled scales (Fig. 30).




Fig. 30

DENTITION: Fourth mandibular teeth received in an internal socket in the upper jaw.Mandibular symphysis to the fourth or fifth tooth. Teeth often tipped with orange,posteriorly rooted in a common elongated socket.Formula: 4 + 14-15

21-22

SIMILAR SPECIES: See P. palpebrosus. Genus Alligator : has more than 25 transverseventral rows; five teeth on each side of the pre-maxillary. Paleosuchus palpebrosus: twotransverse rows of post-occipitals, dorsals reduced to four scales over the hind limbs;ear coverlets not contrasting to the cranial table; no dark median stripe on the dorsalsurface of the snout.

DISTRIBUTION: South America. From Bahia, Brazil, in the south; west to northernBolivia and the tributaries of the Amazon river in eastern Peru; north through Colombiato Ciudad Bolivar on the Rio Orinoco, Venezuela; eastward through Guiana, Surinam,and Mexiana Island

COMMENTS: See Paleosuchus palpebrosus.

REFERENCES: Boulenger, 1889; DeCarvalho, 1955; Medem, 1952, 1958b, 1967;

Mook, 1921b; Schmidt, 1928

a; Schneider, 1801; Wermuth, 1953; Wermuth and

Mertens, 1961; Werner, 1933.




Family: Crocodylidae

Crocody lus acutus CuvierAmerican crocodileCONTENT: No subspecies are recognized.

SIZE: Maximum size of 700cm. suggested. Average adult, 300cm to 400cm. Hatchlings,25cm to 30cm.

COLORATION: Adults gray-green, dark olive-green, or gray-brown above with darkcross-bands on the back and tail, obscure in the old adult. Ventrals white oryellow-white. Lateral ventral scales spotted with gray or black. Sub-caudal scale rowsdensely spotted with gray or black. Young colored as the adult, although with a basecolor of gray rather than green. Individuals from Cuba are dark olive-green withyellowish flecks above. These individuals appear nearly black- above. The commoncoloration closely resembles, but is darker than, the coloration of Crocodylus rhombifer.

CONFORMATION: Skull and snout low, moderately slender, and elongate. Snoutlength 1.8 to 2.5 times the width at the base. Width of the mandible at the posterior pointof the mandibular symphysis less than the distance from the same point to the tip of thesnout. Adults and young adults develop a prominent median hump-like swelling on thesnout anterior to the orbits when viewed in profile. The snout is otherwise smooth andunornamented. The dorsal scalation is highly irregular and variable among individuals.Many of the scales on the dorsal surfaces of the neck and body are separated fromeach other by soft skin. The skin between the enlarged scales is for the most partsmooth, soft, and without many small raised scales. The dorsal scales are restricted tothe median region of the back and are flanked by individual separated enlarged scales,also found on the sides of the body. ISO’s are present on the head and body scalation.The fingers are webbed.

SCALATION:Post-occipitals: a single transverse row of 4 to 6 enlarged scales medially dividedequally, otherwise usually in contact with each other.Nuchals: 4 enlarged scales in a quadrate, flanked by a scale on each side, togetherforming a cluster greatly separated from the dorsal scales by an expanse of smoothskin. Variability in nuchal pattern is great among individuals and may be represented bya single scale or more than 6 scales. The nuchals are greatly separated from thepost-occipitals.Dorsals: 16 to 17 transverse rows of enlarged scales, I to 6 scales per row.Double crest caudal whorls: 16 to 17.Single crest caudal whorls: 15 to 16.Ventral collar: a single enlarged transverse series of scales.Ventral scales: 26 to 32 transverse rows.Sub-caudal scale rows: uniform uninterrupted rings.

Anterior lateral tail inclusions. Usually has a number of scale inclusions.Flanks: A random assortment of large keeled scales surrounded by soft skin (Fig. 31).




Fig. 31

DENTITION: Fourth mandibular teeth received into an external notch on the sides of theupper jaw. Mandibular symphysis extends to the level of the fourth or fifth teeth. Firsttwo mandibular teeth often piercing the upper jaw in adults. All teeth individuallysocketed. Formula: 5 + 13-14.

15.

SIMILAR SPECIES: Crocodylus intermedius: dorsal color light yellow, yellow-green, orwhitish yellow with dark cross-bands on the back and tail; Mandibular symphysisextends the level of the sixth tooth occasionally reaching the level of the seventh tooth.

Crocodylus moreletii : width of the mandible, at the posterior point of the mandibularsymphysis, equal to or greater than the distance from the same point to the tip of thesnout. Adult size less than 250cm; individuals a meter or more in length lose the darkbars on the back and tail and become nearly uniform dark green or gray. Sub-caudalrings interrupted by irregular groups of small scales. Crocodylus niloticus: nuchal scalesclosely associated to the dorsal scales, separated only by one or two small scales;dorsal scales arranged in uniform rows, extending well onto the sides of the body.

DISTRIBUTION:, Extreme southern coastal Florida and the Florida keys In the UnitedStates. From Mexico, once found as far north as Mazatlan on the Pacific coast. ThroughCentral America to coastal Colombia and Venezuela. In the West Indies, particularly

from Cuba, Hispaniola and Jamaica. Not found in Puerto Rico.

COMMENTS: First reported in Florida in 1822, first collected in 1869. Hybridizes freelyin captivity with Crocodylus rhombifer . Extreme difficulty in separating young and

juveniles of this species from those of Crocodylus moreletii on the west coast of Mexico.Such individuals probably led to the proposed use of Crocodylus mexicanus Dumeriland Bocourt (1870).




REFERENCES: Allen and Neill, 1953; Barbour,1923; Boulenger, 1889; Cope, 1900;

Cuvier, 1807;DeSola,1930, 1933; Medem,1962, 1968,1981,1983; Mook, 1921a,b

;

Moore, 1953;Schmidt, 1924, 1944; Thorbjarnarson, 1992; Varona, 1966;Wermuth,1953; Wermuth and Mertens, 1961; and West, 1900.

Crocodylu s cataphractus Cuvier African slender-snouted crocodileCONTENT: No subspecies are recognized.

SIZE: Maximum size given is 400cm. Large adults today rarely more than 300cm, and200cm to 250cm are average. Hatchlings probably average about 25cm.

COLORATION: Adults dark olive green or brown above, with 10 to 13 dark brown orblack cross-bands on the back and tail. The head is un-patterned, occasionally withdark blotches on the sides of the lower jaw, dorsally dark brown or green. Hatchlings

are more vividly colored, some with dark and yellow flecks. Ventral surfaces usuallyuniform cream or white. Specimens from Liberia are often ventrally patterned withnumerous large black spots, several on the sides of the jaws.

CONFORMATION: The cranial table is square and high; the eyes set high on the skull.The snout is unornamented, slender, smooth, and elongate, noticeably concave. Length2.7 to 3.3 times the width at the base. The fourth mandibular teeth usually protrude wellabove the snout in the adult. The dorsal scales are well keeled. The transverse dorsalscale rows are restricted to the middle of the back and do not extend down onto thesides of the body. The fingers of the front feet are slightly webbed at the base. ISO’s arepresent on the head and body scalation.

SCALATION:Post-occipitals: I to 2 pairs of small scales formed as follows: 2 scales isolated fromeach other, usually with 2 additional scales posterior and diagonal to the first pair ofisolated scales. These are divided equally along the midline by smooth skin bearing 2longitudinal rows of nearly indistinct small scales.Nuchals: the nuchals are preceded by a transverse row of 4 to 6 small scales. Thenuchals are composed of 4 transverse rows of 2 scales each continuous with the dorsalscales.Dorsals: 18 to 19 transverse rows of 6 scales each.Double crest caudal whorls: 16 to 18.

Single crest caudal whorls: 15 to 16.Ventral collar: a single transverse row of enlarged scales.Ventral scales: 25 to 29 transverse rows.Sub-caudal scale rows: uniform uninterrupted rings.Flanks: Several large irregularly arranged separated scales sometimes form a loose rowon the sides of the body (Fig. 32).




Fig. 32

DENTITION: Fourth mandibular teeth exposed in an external notch on the sides of theupper jaw. First mandibular teeth often pierce the tip of the snout in adults. Mandibular

symphysis extending to the level of the seventh or eighth teeth. The teeth are long andneedle-like along the entire jaw. All are rooted in widely separated individual sockets.Formula: 5 (4) + 13-14.

15-16

SIMILAR SPECIES: Crocodylus johnsoni , nuchal scales arranged in a quadrate, flankedby two scales nearly continuous with the dorsal scales. Crocodylus intermedius: nuchalscales arranged in a quadrate, flanked by two scales separated from the post-occipitalsand the dorsal scales. Tomistoma schlegelii . mandibular symphysis extending to thelevel of the fourteenth to fifteenth tooth. Gavialis gangeticus: mandibular symphysisextending to the level of the twenty-third to twenty-fourth tooth.

DISTRIBUTION: West Africa, with a single east African locality at Ujiji, a tributary ofLake Tanganyika. Rivers south of Senegal and as far as the Congo River system.Reported from Fernando Po.

COMMENTS: Tomistoma schlegelii (S. Muller) skull figure reproduced in error asCrocodylus cataphractus, Wermuth and Mertens, 1961. Fuchs (1974) describedCrocodylus cataphractus congicus, subsequently included in Wermuth and Mertens(1977), as subspecies, based on commercial undocumented partial skins.

REFERENCES: Boulenger, 1889; Cuvier, 1824;Mook, 1921b; Schmidt, 1919, 1944;

Wermuth,1953; Wermuth and Mertens, 1961; Werner, 1933.

Crocodylu s intermedius GravesOrinoco crocodileCONTENT: No subspecies recognized.




SIZE: Adults reported to 678cm. Males the larger, females usually less than 400cm. to450cm. Hatchlings, 22cm to 25cm.

COLORATION: Adults light yellow, yellowish green above, yellow laterally. Gray orblack cross-bands on the back and tail. Ventral white or yellowish white, un-patterned to

the cloacal vent. Sub-caudal scale rows with dark gray or black spots. Adults mayappear completely white while basking. Juveniles are patterned as the adults, althoughlack green and dorsally are blotched with irregular dark spots. Melanistic individualsfrom Angostura reported but unconfirmed by Medem.

CONFORMATION: Head and snout elongate and slender. Snout length 2 to 2.5 timesthe width at the base. Only a slight swelling rather than a median pre-orbital hump infront of the eyes on the adult. Snout otherwise unornamented. The dorsal scales arearranged in nearly uniform longitudinal rows and overall demonstrate more of aregularity in arrangement than C. acutus. The broad dorsal scales extend laterally ontothe sides of the body. ISO’s are present on the head and body scalation. The fingers

are webbed.

SCALATION.Post-occipitals: a single transverse row of 4 to 6 enlarged scales equally dividedmedially by smooth skin, otherwise in contact with each other.Nuchals: 4 enlarged scales in a square flanked by a scale on each side, forming acluster, widely separated from the dorsal scales by smooth skin.Dorsals: 16 to. 17 transverse rows of 4 to 6 scales.Double crest caudal whorls: 17 to 19.Single crest caudal whorls: 17 to 18.Ventral collar: a single transverse row of enlarged scales.Ventral scales: 25 to 27 transverse rows of 14 to 18 scales.-Sub-caudal scale rows: uniform uninterrupted rings.Flanks: A longitudinal row of six to eight enlarged scales (some larger than the dorsalscales) on the sides of the body (Fig 33).

Fig. 33




DENTITION: Fourth mandibular teeth received into an external notch on the sides of theupper jaw. Mandibular symphysis extends to the level of the sixth teeth, possibly as faras the seventh teeth. The teeth are elongated and needle-like, each individuallysocketed. Formula: 5 + 14.

15

SIMILAR SPECIES: Crocodylus acutus: dorsal color green, gray-green; mandibularsymphysis extends to the level of the fourth or fifth tooth. Crocodylus cataphractus:nuchals arranged in four transverse rows of two scales, continuous with the dorsalscales. Crocodylus johnsoni : nuchals formed in a cluster of scales, continuous with thedorsal scales.

DISTRIBUTION: Restricted to the Orinoco river system of Venezuela and easternColombia, and occasionally wafted to Trinidad. May have ranged into the Guyana’s atone time.

COMMENTS: The species has been confused with other crocodilians in the literature:the. Crocodylus intermedius skull figured in Mook (1921b), Wermuth (1953), Wermuthand Mertens (1961) is an example of Crocodylus cataphractus Cuvier. Temsacusintermedius Gray (= C. intermedius Graves) in Rochebrune (1883) is erroneouslyincluded in the fauna of Africa. The Orinoco crocodile life figure in DeSola (1933),Ditmars (1913), and Wermuth (1953) is an example of Crocodylus. cataphractus Cuvier.Much of this has been clarified by Brazaitis (1973).

REFERENCES: Boulenger, 1889; Brazaitis,1973; DeSola, 1833; Ditmars, 1913;Graves, 1819; Gray, 1869; Lutkin, 1884 Medem, 1958, 1968; Mook, 1921b,Rochebrune, 1883; Schmidt, 1919; Thorbjarnarson, 1992; Wermuth, 1953; Wermuthand Mertens, 1961; Werner, 1933.

Crocody lus johnson i KrefftJohnston’s crocodile CONTENT: No subspecies are recognized.

SIZE: Average adult 180cm to 240cm. Maximum size 300cm. Hatchlings approximately20cm to 25cm.

COLORATION: Dark brown or olive green above, flecked with black. Some faint darkcross-bands on the back and tail. Ventrals uniform white.

CONFORMATION: Snout slender and elongate, length 2.3 to 3.2 times the width at thebase (Fig 34). Snout smooth and unornamented. The teeth form a perfectly interlockingzigzag pattern along the sides of the jaws. The dorsal scales are formed in nearlyperfect length rows, as are the longitudinal rows of scales, which line the sides of thebody. The fingers are not webbed. ISO’s are present on the head and body scalation.




SCALATION:Post-occipitals: a single transverse row of 4 enlarged scales, separated medially butotherwise in contact with each other.Nuchals: 4 scales in a square, flanked by a scale on each side, forming a cluster.Nuchal scales continuous with the dorsal scales (Fig 34a).

Dorsals: 18 transverse rows of 6 to 8 scales.Double crest caudal whorls: 17.Single crest caudal whorls: 16 to 17.Ventral collar: a single transverse row of enlarged scales.Ventral scales: 22 to 24 transverse rows.Sub-caudal scale rows: The majority of individuals bear rows that are interrupted byseveral irregular groups of small scales.Flanks: Uniform series of unremarkable scales. (Fig. 34b).

DENTITION: Fourth mandibular teeth exposed in an external notch on the sides of the

upper jaw. Mandibular symphysis extends to the level of the sixth or eighth teeth. Thefirst mandibular teeth may pierce the snout. Formula: 5+ 14-16.15

SIMILAR SPECIES: Crocodylus intermedius; Crocodylus acutus: nuchal scales widelyseparated from the dorsal scales by an expanse of skin. Crocodylus cataphractus:nuchals are arranged in four transverse rows of two scales each, continuous with thedorsal scales.

Fig. 34

Fig. 34b

Fig. 34a




DISTRIBUTION: Northern Australia, from the Fitzroy River in the west to about latitude210 south in the east.

COMMENTS: The spelling of the common name has been the subject of some

confusion. The original type was collected by sub-Inspector Johnstone in the HerbertRiver in Queensland. However, the collector's name was misspelled by Gray (1873) indescribing the type as Crocodylus johnsoni .

REFERENCES: Barrett, 1950; Boulenger, 1889; Gray, 1874; Krefft, 1873; Wermuth,1953; Wermuth and Mertens, 1961; Werner, 1933; Worrell, 1963.

Crocody lus mind orens is Schmidt Philippine crocodileCONTENT: No sub-species are recognized

SIZE: Maximum size up to 250cm; average 150cm. to 200cm.

COLORATION: Dorsal color brown, transverse dorsal bars on the back and tail. Shortdark bars extend onto the ventral scales on the sides. Ventrals are white.

CONFORMATION: Snout moderate, length 1.8 to 2.3 times the width at the base. Tworidges extend well onto the snout from the anterior points of the orbits (Fig 35). Transverse dorsal rows broad at mid-body extending laterally onto the sides. One ortwo irregularly arranged longitudinal rows of three to six enlarged scales on each side ofthe body (Fig 35a) . Fingers are slightly webbed at the base. ISO’s are present on thehead and body scalation.

SCALATION:Post-occipitals: a single transverse row of 4 to 6 enlarged scales, separated medially,but otherwise in contact with each other.Nuchals: 4 enlarged scales in square, flanked by a scale on each side, forming acluster. Nuchals closely associated to but separated from the dorsals by two smallscales. Scales making up the cluster are in contact with each other and are notseparated along the midline by soft skin.Dorsals: 17 transverse rows of 6 to 8 scales.Double crest caudal whorls: 18 to 19.Single crest caudal whorls: 16 to 18.Ventral collar: a single transverse row of enlarged scales.Ventral scales: 25 to 26 transverse rows.Sub-caudal scale rows: uniform uninterrupted rings.Flanks: Similar to Crocodylus ng. novaeguineae. (See Fig. 38)




Fig 35 Fig 35a

DENTITION: Fourth mandibular teeth exposed in an external notch on the sides of theupper jaw. Mandibular symphysis extends to the level of the fourth or fifth teeth. Allteeth are independently socketed. Formula: 5 + 13-14.

15

SIMILAR SPECIES: Crocodilus novaeguineae: nuchal cluster separated along themidline by soft skin; scales on the sides of the body uniform in size, arranged in uniformlongitudinal rows. Crocodylus porosus: lacks post-occipital scales or displays only asingle pair of small scales; Crocodylus siamensis: Series of irregular scales interrupt themedial anterior sub-caudal scales; more than 29 transverse ventral rows.

DISTRIBUTION: The Philippine Islands of Luzon, Mindoro, and Mindanao

COMMENTS: The ecological relationship between this species and Crocodylus porosus is not clearly understood, nor is the parallel relationship to the Philippine race. Wermuth(1953) placed the New Guinea and Philippine crocodiles in species synonymy. DistinctVariations among populations, leading to the separation of the two taxa into separatespecies were reported by Ross (1983, 1986 unedited, unreviewed) and Hall (1989)

REFERENCES: Riosuke, 1985; Ross, 1986; Ross and Alcala, 1983;Schmidt,1935,1938,1944,1956; Wermuth,1953; Wermuth and Mertens, 1961; 1977;Werner, 1933.




Crocodylu s morelet ii Dumeril and Bibron Morelet’s crocodile CONTENT: No subspecies are recognized.

SIZE: Adults rarely 250cm. Average size 100cm to 150cm. Hatchlings, 22cm to 27cm.

COLORATION: Young are dark olive green with dark cross-bands on the back and tail.Some yellow flecks on the dorsal surfaces and yellow irregular striations on the sides.Overall becomes darker with age, nearly uniform dark green or black at about I00cm.Some dark irregular striations extend onto the lateral ventral surfaces in the form ofshort bars.

CONFORMATION: The head is flat and wide, the eyes small and set low into the skull.The -snout is massive and wide, length 1.5 times the width at the base. A median ridge,slightly humped anterior to the orbits extends down well onto the snout. The

squamosals are nearly lost in the flattened, laterally rounded, cranial table. The scalesof the back, sides, and tail are feebly keeled. The flanks are smooth. The transverserows of dorsal scales are flanked by irregularly arranged separated scales on the backand sides of the body (Fig. 36). ISO’s are present on the head and body scalation. Thefingers are webbed.

SCALATION:Post-occipitals: a single transverse row of 4 to 6 enlarged scales equally dividedmedially by smooth skin, each scale separate from each other.Nuchals: 4 enlarged scales in a square flanked by a scale on each side, forming acluster, separated from the dorsals by a space of soft skin.Dorsals: 15 to 17 transverse rows of 4 to 6 scales in contact with each other and flankedby irregularly arranged separated scales.Double crest caudal whorls: 19 to 20 (17 to 18 reported).Single crest caudal whorls: 19.Ventral collar: a single transverse row of enlarged scales.Ventral scales: 29 to 33 transverse rows of sub-caudals. The rings are interrupted by anumber of irregular series of scales.Flanks: One or two rows of enlarged scales. Upper rows surrounded by grainy skin,lower rows surrounded by distinct scales.(Fig. 36a).




DENTITION: Fourth, mandibular teeth exposed in an external notch on the sides of theupper jaw. First mandibular teeth often pierce the upper jaw. Mandibular symphysisextends to the level of the fourth or fifth teeth. Mandible wider at the posterior point ofthe symphysis than from the same point to the tip of the jaw. Teeth are rather short andthick,. rooted in individual sockets. Formula: 5 + 13-14.

15

SIMILAR SPECIES: Crocodylus acutus: dorsal color green, gray-green with dark cross-bands on the back and tail, Sub-caudal scale rows in uniform uninterrupted rings.

Crocodylus rhombifer : toes short and not webbed; Sub-caudal scale rows in uniformuninterrupted rings. Crocodylus palustris: color brown or yellow-brown with dark cross-bands; Sub-caudal scale rows in uniform uninterrupted rings. Crocodylus n.novaeguineae: less than 26 transverse ventral rows. Crocodylus niloticus: from 26 to 30transverse ventral rows; nuchals in close association to the dorsal rows.

Fig. 36

Fig. 36a




DISTRIBUTION: Pacific coast of Mexico from the state of Nayarit and Sinaloa south. Atlantic coast from Tamaulipas south through Yucatan, British Honduras, andGuatemala

COMMENTS: Young animals easily confused with the young of Crocodylus acutus on

the west coast of Mexico. The species was not recognized by Mook (1921) as distinctfrom C. rhombifer , a most closely related species, ecologically.

REFERENCES:. Boulenger, 1889; Dumeril and Dumeril, 1851; Mook, 1921b; Powell,

1965; Schmidt, 1924; Thorbjarnarson, 1992; Wermuth, 1953; Wermuth and Mertens,1961; Werner, 1933.

Crocody lus n i lo t icus LaurentiNile crocodileCONTENT: No subspecies are currently recognized.

SIZE: Adults 700cm would be extremely large individuals. Average size would be400cm to 600cm. Hatchlings, 26cm to 34cm.

COLORATION: Adults and young dark olive green, yellow-green, brown with darkcross-bands on the back and tail. Old adults uniform dark brown or green with faint darkcross-bands. Dark spots and irregular striations on the sides of the body extends downonto the lateral ventral scales. Ventrals are white, light gray, or cream. Specimens fromMadagascar are densely flecked with 6rk brown or black on the head, back, and tail.

CONFORMATION: The snout is moderate, extremely rugose, length 1.6 to 2 times the

width at the base. The transverse dorsal scales are broad at the mid-body and extendonto the sides. The lateral body scales are large, barely discernible from the dorsalscales in size but feebly keeled. ISO’s are present on the head and body scalation. Thefingers are slightly webbed at the base.

SCALATION:Post-occipitals: a single transverse row of 4 to 6 enlarged scales divided equally at themidline, otherwise in contact with each other.Nuchals: 4 large scales in a square, flanked by a scale on each side, forming a cluster.Nuchals are separated from the dorsal scales by I to 2 small scales.Dorsals: 17 to 18 transverse rows of 6 to 8 small scales.

Double crest caudal whorls: 17 to 18.Single crest caudal whorls: 15 to 16.Ventral collar: I to 2 transverse rows of slightly enlarged scales.Ventral scales: 26 to 30 transverse rows.Sub-caudal scale rows: usually uniform uninterrupted rings.Flanks: Usually only a one or two rows of enlarged scales on the sides of the body.(Fig 37)




Fig. 37

DENTITION: Fourth mandibular tooth exposed in an external notch on the sides of theupper jaw. Mandibular symphysis extending to the level of the fourth or fifth teeth. Allteeth are individually socketed. Formula: 5 + 13-14.

14-15

SIMILAR SPECIES: Crocodylus p. palustris, Crocodylus p. kimbula: some transversedorsal rows reduced to four scales per row; ventral collar consists of a single transverserow of enlarged scales; some populations without a noticeable ventral collar.Crocodylus novaeguineae, Crocodylus mindorensis: ventral scales in 24 to 26transverse rows. Crocodylus siamensis: from 30 to 34 transverse ventral rows; alongitudinal bony ridge raised between the orbits on the frontal region of the skull.

DISTRIBUTION: All of Africa, below the Sahara. Found in Madagascar, the Comores,

and once reported from the Seychelles Islands. No longer found in the wild in Israel.

COMMENTS: Some populations have well developed dermal ossifications in the ventralregions. Fuchs (1974) described Crocodylus niloticus africanus, as a subspecies fromTanzania, Ruanda, Urundi; C. n. chamses as a subspecies from Uganda, southeasternSudan, Republic of Congo, Zaire, Gabon, Angola, northern southwest Africa andnortheast Sambia; C. n. cowei, from Mozambique, Botswanaland, Malawi, Rhodesia,Sambia, South Africa, and Sudan; C. n. madagascariensis from Madagascar, theKomore and Sechelle Islands; C. n. pauciscutatus from Kenya, Ethiopia, the SeychelleIslands; and C. n. suchus from Nigeria, Niger, Cameroon, Chad, Central AfricanRepublic, Equatorial Guinea, Senegal, Mali, Guinea, Gambia, Volta, Ghana, Togo,

Dahomey, subsequently included in Wermuth and Mertens (1977), as subspecies,based on commercial undocumented partial skins.

REFERENCES: Boulenger, 1889; Cott, 1961;Grandidier, 1872; Laurenti, 1768; Modha,1967; Mook, 1921b; Pooley, 1966; Schmidt, 1919, 1944; Thorbjarnarson, 1992;Wermuth, 1953; Wermuth and Mertens, 1961




Crocodylu s nov aeguin eae Schmidt New Guinea crocodileCONTENT: No sub-species are recognized.

SIZE: Maximum size 284cm. Average adult is 180cm to 200cm. Hatchlingsapproximately 25cm.

COLORATION: Drab olive green, gray-green above with interrupted dark cross-bandson the back and tail; extending onto the sides but not onto the ventral scales. Dorsalsurfaces flecked with dark brown or black. Ventrals uniform white.

CONFORMATION: Snout elongate, length 1.8 to 2.3 times the width at the base. Aslight suggestion of a pre-orbital ridge present at the anterior points of the orbits in adultindividuals. The dorsals are broad at the mid-body, extending onto the sides. Thefingers are webbed. ISO’s are present on the head and body scalation.

SCALATION:Post-occipitals: a transverse row of 4 enlarged scales, divided equally at the midline,separated from each other.Nuchals: variable in size and arrangement. Four scales in a square, with a scale oneach side, forming a cluster. Nuchal scales tend to be separated from each other,particularly along the midline; which divides the cluster with a space of smooth skin.Nuchals separated from the dorsals by 2 small scales only.Dorsals: 17 transverse rows of scales. The widest at mid-body bears 10 scales.Double crest caudal whorls: 18 to 20.Single crest caudal whorls: 17 to 19.Ventral collar: a single transverse row of enlarged scales.Ventral scales: 24 to 25 transverse rows of scales.Sub-caudal scale rows: uniform uninterrupted rings.Flanks: Flank scales are oval, arranged in uniform longitudinal rows alternating withrows of smaller oval scales (Fig. 38).

Fig. 38





15

SIMILAR SPECIES: Crocodylus mindorensis: sides of the body bear one or two slightlyirregular rows of enlarged scales; nuchal scales in contact with each other, notseparated along the midline. Crocodylus porosus: Ventrals in more than 29 transverserows; post-occipitals usually absent. Crocodylus siamensis: a longitudinal raised ridgebetween the orbits on the frontal; ventrals in more than 30 transverse rows; a medianrow of irregular scales extending from the vent through the first several rows of sub-caudal whorls.

DISTRIBUTION: Inland, New Guinea

COMMENTS: The ecological relationship between this species and Crocodylus porosus

is not clearly understood, nor is the parallel relationship to the Philippine race. Wermuth(1953) placed the New Guinea and Philippine crocodiles in species synonymy. DistinctVariations among populations, leading to the separation of the two taxa into separatespecies were reported by Ross (1986 unedited, un-reviewed) and Hall (1989)

REFERENCES: Hall, 1989; Neill, 1946; Ross,1986; Schmidt, 1928b, 1932, 1944; Smith,

1931; Wermuth, 1953; Wermuth and Mertens, 1961; Werner 1933.

Crocodylu s palustr is palustr is LessonMugger, Marsh crocodile, Indian crocodile

CONTENT: Crocodylus palustris is composed of two races: Crocodylus palustris palustris and Crocodylus palustris kimbula.

SIZE: Adults recorded up to 400cm. Average adult 200cm to 300cm. Hatchlingsapproximately 27cm.

COLORATION: Brown, yellow-brown, or olive-brown, with dark cross-bands on the backand the tail. Dark irregular striations on the sides of the body and tail barely enter theventral scales. Ventrals uniform white.

CONFORMATION: Head and snout massive and rugose. Eyes set low into the skull.

Snout length 1.3 to 1.5 times the width at the base. The dorsal scales are restricted tothe median dorsal region and do not extend down onto the sides of the body. Twolongitudinal rows of five to six enlarged scales on each side of the body. The fingers areslightly webbed at the base. ISO’s are present on the head and body scalation.

SCALATION:Post-occipitals: usually a single transverse row of 4 to 6 enlarged scales, separatedequally at the midline, but otherwise in contact with each other.




Nuchals: 4 scales in a square, flanked by a scale on each side, forming a cluster;closely associated to but separated from the dorsals by two small scales.Dorsals: 16 to 18 transverse rows of 4 to 6 scales. The majority of scale rows contain 4scales.Double crest caudal whorls: 18 to 19.

Single crest caudal whorls: 16.Ventral collar: no distinct transverse series of enlarged scales.Ventral scales: 26 to 32 transverse rows.Sub-caudal scale rows: uniform uninterrupted rings.Flanks: Upper flank scales surrounded by bead like scales, lower scales surrounded bydistinct large scales (Fig. 39).

Fig. 39

DENTITION: Fourth mandibular teeth exposed in an external notch on the sides of theupper jaw. Mandibular symphysis extends to the level of the fourth or fifth teeth. All

teeth are independently socketed. Formula: (4) 5 + 14.15SIMILAR SPECIES: Crocodylus p. kimbula: ventral collar present as a single transverseseries of enlarged scales. Crocodylus moreletii : Sub-caudal scale rows interrupted byirregular groups of small scales; dorsal color dark olive green or black. Crocodylus

porosus: post-occipitals usually absent, or a single pair of slightly enlarged separatedscales; two pre-orbital ridges extend well onto the snout from the anterior points of theorbits.

DISTRIBUTION: India and West Pakistan to the Brahmaputra of west Assam, to Iran. Asingle report from Burma is doubtful. Two specimens of questionable locality, "ThePhilippines," were examined (AMNH 4805 and AMNH 4806).

COMMENTS: The validity of the two races that comprise the species is somewhatquestionable. Although there are relatively minor morphological differences separatingthe two races, there appear to be substantial behavioral and ecological separations.




REFERENCES: Boulenger, 1889; Deraniyagala, 1936, 1939; DeRooij, 1915; Loveridge,

1945; Mook, 1921b; Schmidt, 1944; Smith, 1928, 1931; Wermuth, 1953; Wermuth and

Mertens, 1961; Werner, 1933; Yadov, 1968.

Ceylon Marsh crocodile Crocody lus palus t r is k imb ula Deraniyagala CONTENT: Crocodylus palustris is composed of two races: Crocodylus palustris

palustris and Crocodylus palustris kimbula.

SIZE: Adults 350cm to 400cm; record is 557cm. Average size about 200cm, in recenttimes. Hatchlings 25cm-to 27cm.

COLORATION: As Crocodylus p. palustris.

CONFORMATION: As in Crocodylus p. palustris.

SCALATION:Post-occipitals: as in Crocodylus p. palustris.Nuchals: as in Crocodylus p. palustris.Dorsals: 16 to 18 transverse rows of 4 to 6 scales. The majority of transverse rowscontain 6 scales.Double crest caudal whorls: 18 to 19.Single crest caudal whorls: 16.Ventral collar: a single transverse row of enlarged scales.Ventral scales: 26 to 32 transverse rows.

Sub-caudal scale rows: uniform uninterrupted rings.Flanks: See Fig. 39.

DENTITION: As Crocodylus p. palustris. Formula: (4) 5 + 14.15

SIMILAR SPECIES: Crocodylus p. palustris: ventral collar absent, composed of scaleswhich are not noticeable larger than the ventral scales. Additional as C. p. palustris.

DISTRIBUTION: Lowland areas and plains of Sri Lanka

COMMENTS.: As Crocodylus p. palustris.

REFERENCES: Boulenger, 1889; Deraniyagala, 1936,1939; DeRooij, 1915; Loveridge,

1945; Mook, 1921b; Schmidt, 1944; Smith, 1928, 1931; Wermuth, 1953; Wermuth and

Mertens, 1961; Werner, 1933; Yadov, 1968.




Crocody lus porosus Schneider Saltwater crocodile, estuarine crocodile, Indo-Pacific crocodileCONTENT: No subspecies are recognized.

SIZE: The largest reptile and crocodilian in the world today. Maximum recorded size is

1005 cm (collected in Bengal in 1840). Average adult, 350 cm to 450 cm; hatchlings, 22cm to 30 cm.

COLORATION: The hatchlings and young adults are yellow above with numerous largedark spots on the head, body, and tail. Adults become dark olive drab above with slightyellow remaining on the sides. Some of the dark spotting is also retained. The ventralscales are immaculate white.

CONFORMATION: Snout moderate, becoming massive with age, rugose, length 1.7 to2.1 times the width at the base. Two longitudinal bony ridges extend onto the snout fromthe anterior points of the orbits. Squamosals are raised into a bony ridge. Dorsal scales

are restricted to the median portions of the back and do not extend laterally onto thesides of the body.. The fingers are webbed. ISO’s are present on the head and bodyscalation.SCALATION:Post-occipitals: absent entirely or represented by 1, rarely 2, pairs of small slightlyenlarged, well separated, scales. The skin in the post-occipital region is composed ofnumerous, irregularly arranged, small bead-like scales.Nuchals: 4 large scales arranged in a square, with a scale on each side forming acluster widely separated from the dorsal scales.Dorsals: 16 to 17 transverse rows of 6 to 8 scales.Double crest caudal whorls: 19 to 20.Single crest caudal whorls: 19 to 21.Ventral collar: a single enlarged row of transverse scales.Ventral scales: 31 to 35 transverse rows.Sub-caudal scale rows: some individuals have rings interrupted by irregularly arrangedsmall scales.Flanks: Scales of the sides of the body are small, uniform in size, and arranged inuniform longitudinal rows (Fig. 40).

Fig. 40




DENTITION: Fourth mandibular teeth exposed in an external notch on the sides of theupper jaw. Mandibular symphysis extends to the level of the fourth or fifth teeth. Allteeth are independently socketed. Formula: 4 (5) + 13-14.

15

SIMILAR SPECIES: Crocodylus siamensis: a series of irregular scales extendingmedially from the vent through the first several rows of sub-caudal scale rows; a slightlyraised bony ridge extends from the anterior point of the orbit and extends for only ashort distance onto the snout; from two to four enlarged post-occipital scales.Crocodylus novaeguineae and Crocodylus mindorensis: two to three pairs of welldeveloped post-occipital scales, ventral scales in less than 28 transverse ventral rows.

DISTRIBUTION: Australia, southern coastal China, throughout Southeast Asia,Indonesia, Papua New Guinea, the Indo Pacific Islands of the Solomon’s, Republic ofPalau, and Vanuatu; historically Singapore and Hong Kong. Gulf coast of India and Sri

Lanka.

COMMENTS: Crocodylus porosus minikanna was described as a subspecies occurringin Sri Lanka. It is not considered here. Crocodylus porosus life figure in Barret (1956) isan example of Alligator mississippiensis. Australian C. porosus are probably the mostcompletely investigated species of crocodilian, based primarily on the extensive worksof H. Messell and et al. (1978-1987, 20 monographs), and Grahame Webb, WildlifeManagement Int. Pty. Ltd., Sanderson, NT, Australia. Fuchs (1974) restrictedCrocodylus porosus porosus to Sri Lanka, and defined Crocodylus porosus biporcatusas the subspecies occurring throughout the remainder of the species range, and

subsequently included the taxon in Wermuth and Mertens (1977).

REFERENCES: Barrett, 1950; Boulenger, 1889;DeRooij,1915; Kimura, 1968;

Loveridge, 1945; Mook,1921b; Schmidt, 1932, 1944, 1956;Schneider, 1801; Smith,

1931; Wermuth, 1953;Wermuth and Mertens 1961; Messell et al., 1978-1987; Werner,1933; Worrel, 1963.

Crocody lus rhom bi fer CuvierCuban crocodileCONTENT: No subspecies are recognized.

SIZE: Adults rarely more than 350 cm. A maximum size of 500 cm is suggested. Average adult 200 cm to 250 cm. Females larger than males in average size, reported.Hatchlings little known, approximately 20 cm to 30 cm.

COLORATION: Adults yellow-green or green above with dense dark green or black.speckling on the head, body, and tail. The post-occipitals, nuchal, and dorsal scales areoften extremely dark or black. The small raised pointed scales which cover the skinbetween the large body scales are tipped with yellow and black. The limbs, sides of




the body, and tail are blotched with yellow and black irregular striations. A series ofdark bars extends onto the ventral scales from the side irregular striations. Ventralsurfaces are white, laterally tinged with yellow on some individuals. The edges of theeyelids are usually white, giving a "spectacled" appearance. Old adults become darkerwith age but retain the yellow and black blotches and irregular striations on the sides of

the body and tail. Juveniles are patterned as the adults in vividly rich tones.

CONFORMATION: The head and jaws are massive. The snout rugose, length 1.4 to 1.6times the width at the base. Adults bear a longitudinal ridge on the snout, originating ata point medially anterior to the orbits (suggestive of a pre-orbital hump at this point), andextending down the middle of the snout to the level of the external lateral pre-maxillary/maxillary notches. The squamosals are elevated into a high bony ridge. The extremelyuniformly arranged dorsal scales are confined to the median region of the back and donot extend down onto the sides of the body.. ISO’s are present on the head and bodyscalation. The fingers are extremely short and lack webs.

SCALATION:Post-occipitals: a single transverse row of 4 sharply keeled, enlarged scales, wellseparated from each other. The greatest separation occurring at the midline, dividingthe scales equally. Surrounding scalation is raised and bead-like.Nuchals: 4 enlarged scales in a quadrate flanked by a scale on each side, forming acluster in close association to the dorsal scales but separated from them by I to 2 smallscales.Dorsals: 16 to 17 transverse rows of 6 scales each. Median longitudinal rows onlyfeebly keeled.Double crest caudal whorls: 17 to 19 rings.Single crest caudal whorls: 17 to 18Ventral collar: a single transverse row of enlarged scales.Ventral scales: 32 to 33 transverse rows. The fingers are slightly webbed at longitudinalrows of enlarged scales on each side of the body.Subcaudal scales: uniform uninterrupted rings.the base. ISO’s are present on the headand body scalation.Flanks: There are two to three longitudinal rows of enlarged isolated scales on eachside of the body. Upper flank scales surrounded by bead like scales, lower scalessurrounded by distinct large scales (Fig.41).

Fig.41




DENTITION: Fourth mandibular teeth exposed in an external notch in the sides of theMandibular symphysis extends to the level of the fourth or fifth teeth. All teethindependently socketed. Formula: 5 + 13-14.

15

SIMILAR SPECIES: Crocodylus moreletii : fingers are long and webbed; Sub-caudalscale rows arranged in rings interrupted by groups of irregular small scales; neckscales, keeled but not tubercular in elevation. Crocodylus niloticus, Crocodylusnovaeguineae: ventrals in less than 30 transverse rows. Crocodylus palustris: dorsalscales arranged in irregular rows, the broadest transverse rows extending onto thesides; sub-caudal scale rows, arranged in uniform uninterrupted rings, dorsal color isbrown. Crocodylus acutus: fingers long and webbed; dorsal scales irregularly arranged;snout elongate and slender; Crocodylus siamensis: a series of irregular scales extendmedially from the vent through the first several rows of sub-caudal scale rows, fingerslong and webbed.

DISTRIBUTION: Historically the Isle of Pines, the Zapata swamp, central Cuba

COMMENTS: Hybridization under captive breeding programs, between Crocodylusacutus and Crocodylus rhombifer , threatens the continued genetic integrity of thisspecies

REFERENCES: Barbour and Ramsden, 1919;Boulenger, 1889; Buide, 1967; Cuvier

1807; DeSola, 1930; Mook,1921b; Thorbjarnarson, 1992; Varona, 1966; Wermuth,

1953; Wermuth and Mertens, 1961; Werner, 1933.

Crocodylu s siamensis Schneider Siamese crocodileCONTENT: No subspecies are recognized.

SIZE: Adults 350 cm to 400 cm. Average adult 250 cm to 300 cm. Hatchlings,approximately 25 cm.

COLORATION: Dark olive, yellow, or brownish-green, spotted with black above. Darkcross-bands on the back and tail. Ventrals white or yellowish-white.

CONFORMATION: Snout is moderate and slightly concave, length 1.5 to 1.6 times thewidth of the base. A short pair of sharply converging ridges at the anterior point of theorbits. A longitudinal bony ridge is evident between the orbits on the frontal. Thesquamosals are raised in a high ridge. The dorsal scale rows are broadest at mid-bodyand extend onto the sides. Two irregular




SCALATION:Nuchals: 4 scales in a square, flanked by a scale on each side, forming a cluster; widelyseparated from the post-occipitals as well as the dorsal scales.Dorsals: 16 to 18 transverse rows of 6 scales.Double crest caudal whorls: 19 to 20.

Single crest caudal whorls: 17 to 19.Ventral collar: a single transverse row of enlarged scales.Ventral scales: 30 to 34 transverse rows.Sub-caudal scale rows: anterior several rings interrupted by irregular groups of smallscales. A series of irregular scales extend medially from the vent through the firstseveral rows of sub-caudal scale rows.Flanks: One or two rows of enlarged keeled scales surrounded by small scales(Fig. 42).

Fig. 42


15SIMILAR SPECIES: Crocodylus porosus: lacks a longitudinal ridge between the orbits;yellow or yellow-green above with black spots and cross-bands; . Crocodylus p.

palustris; Crocodylus p. kimbula: nuchal scales closely associated to the dorsal scales,separated only by two small scales. Crocodylus moreletii : no series of irregular scalesextending medially from the vent through the first several rows of sub-caudal scalerows, squamosals flattened, laterally rounded; a slight hump or swelling medially,anterior to the orbits, continuing as a slight ridge down the middle of the snout; thefingers are webbed. Crocodylus novaeguineae and C. mindorensis: less than 28 ventralrows.




DISTRIBUTION: Historically central and peninsular Thailand; Cambodia, the river Quae(north to latitude 161); and the northern part of the Malay peninsula. Questionablereports from Java. Some wild populations may still occur in the Mekong river system.

COMMENTS: Has been bred in large numbers under captive conditions in Thailand.

Wild populations are largely depleted.

REFERENCES: Boulenger, 1889; Cox et al., 1993; DeRooij, 1915; Loveridge, 1945;Schmidt, 1944; Schneider, 1801; Y. Siah in lit.; Smith, 1931; Taylor, 1970; Wermuth,1953; Wermuth and Mertens, 1961; Werner, 1933.

Osteolaemus tetraspis tetraspis CopeWest African dwarf crocodileCONTENT: Osteolaemus tetraspis is comprised of two races: Osteolaemus t. tetraspis and Osteolaemus t. osborni .

SIZE: Adults rarely 200 cm in length. Average size 100 cm to 150 cm. Hatchlings 20-25cm.

COLORATION: Adults uniform black above, blotched or uniform black on the ventralsurfaces. Young are blotched with brown above with dark cross-bands on back and tail.Some yellow irregular striations on the sides of the jaws, body, and tail. Ventral surfacesblotched with brown or black. The iris of the eyes of the young and adults is dark brown.The palate white or cream, anteriorly mottled with black.

CONFORMATION: Snout short, wide and blunt, concave. The nasal region is wellelevated on the snout in the adult. Snout only slightly longer than its width at the base,rugose but without other ornamentation. The eyes are large, eyelids rugose, withwell-developed "eyelash-like" appendages. The dorsal scales are arranged in uniformlongitudinal rows. The two vertebral rows lack any significant keel. ISO’s are present onthe body and head, but difficult to see on the coarse skin of the body. The fingers areslightly webbed at the base. The body is heavily armored, particularly in the ventralregions. Old animals become black with coarse bark-like skin.

SCALATION:Post-occipitals: usually a single transverse row of sharply raised scales.Nuchals: 4 large scales arranged in a square, closely associated to, but separated fromthe dorsals by 2 small scales.Dorsals: 17 to 20 transverse rows of 6 to 8 scales.Double crest caudal whorls: average I 1 (10 to 12).Single crest caudal whorls: 15 to 17.Ventral collar: a single transverse row of enlarged scales.Ventral scales: 26 to 27 transverse rows.Sub-caudal scale rows: uniform uninterrupted rings.




Flanks: The flanks have a number of randomly occurring enlarged keeled and pointedscales. The surrounding skin is pebble-like and coarsely creased (Fig. 43).

Fig. 43

DENTITION: Fourth mandibular teeth exposed in a notch on the sides of the upper jaw.Mandibular symphysis extends to the level of the fourth or fifth teeth. The teeth are shortand thick, individually socketed. Formula: 4 + 12-13

14-15

SIMILAR SPECIES: Alligator sinensis: fourth mandibular teeth socketed; there are fiveteeth on each side of the pre-maxillary. Paleosuchus trigonatus, Paleosuchus

palpebrosus: more than 20 teeth on each side of the lower jaw. 0steoldemus tetraspis

osborni : snout flat, not concave; nostrils not elevated above the snout; interior of themouth (palate) yellowish-white.

DISTRIBUTION: West Africa, south of the Sahara and northwest of the Congo Riversystem. Distributional separation between the two races of Osteolaemus is not clearlydefined.

COMMENTS: See Osteolaemus tetraspis osborni.

REFERENCES: Boulenger, 1889; Cope, 1861; Inger, 1948; Kofron and Steiner,1994;Schmidt, 1919,1944; Wermuth, 1953; Wermuth and Mertens. 1961, Werner, 1933.

Osteolaemus tetraspis osborni Congo dwarf crocodileCONTENT: See Osteolaemus t. tetraspis.

SIZE: Largest specimen collected by Schmidt was 114 cm. Maximum size probably 150cm.




COLORATION: See Osteolaemus t. tetraspis. Some adults display faint, wide browncross-bands on the back and tail. The interior of the mouth (palate) yellowish-white orwhite.

CONFORMATION: See Osteolaemus t.. tetraspis. Snout is flat, not concave. Nasalregion is not elevated. A lateral longitudinal row of approximately five enlarged scaleson each side of the body.

SCALATION:Post-occipitals: 2 transverse rows of-4 scales, separated from each other.Nuchals: 3 pairs of scales arranged in 2 longitudinal rows, ending in close proximity tobut not in contact with the dorsal scales.Dorsals: 18 transverse rows, the broadest containing 16 scales (rows of 4 scalesfollowed by rows of 6 scales, followed by rows of 4 scales).Double crest caudal whorls: 12 to 14.

Single crest caudal whorls: 17 to 19.Ventral collar: a single transverse row of enlarged scales.Ventral scales: 21 to 23 transverse rows.Sub-caudal scale rows: uniform uninterrupted rings.Flanks: See Fig 43.

DENTITION: See Osteolaemus t. tetraspis. Formula: 4 + 12-13.24-15

SIMILAR SPECIES: Osteolaemus t. tetraspis: snout concave, nostrils elevated; interiorof mouth (palate) darkly mottled anteriorly, posteriorly cream or white.

DISTRIBUTION: Northern Congo River system

COMMENTS: Schmidt originally designated this race as a new genus, Osteoblepharon,which has not been upheld.

REFERENCES: Inger, 1948; Schmidt, 1919,1944; Wermuth, 1953; Wermuth andMertens, 1961; Werner, 1933.

Family: Gavialidae

Gavial is g anget icus (Gmelin) GharialCONTENT: No subspecies are recognized.

SIZE: Maximum recorded size 625 cm. Average adult 350 cm to 450 cm. Hatchlingsapproximately 37 cm.




Fig. 44

Fig. 44a

COLORATION: Dark or light olive above with dark cross-bands and speckling on thehead, body, and tail. Dorsal surfaces becoming dark almost gray-black at about 20years of age. Ventrals are yellowish-white.

CONFORMATION: Snout extremely long and slender, parallel sided. Squamosalsrounded and flat. Snout length 3.3 to 5.5 (in young) times the width of the base (Fig.44), smooth and completely unornamented. Males develop a hollow nasal protuberanceat sexual maturity. The neck is elongated and thick. The dorsals are more or lessrestricted to the median regions of the back. The fingers are extremely short and thicklyemarginated with a web. The ventral scales have ISO’s, at least in the gular region.

SCALATION:Post-occipitals: a single enlarged pair of separated scales or two small scales with twoadditional scales at diagonals to the first pair.Nuchals: 4 transverse rows of 2 scales continuous with the dorsal scales.

Dorsals: 21 to 22 transverse rows of 6 scales. (Nuchals + dorsals = 21 to 24 rows).Double crest caudal whorls: 18 to 26.Single crest caudal whorls: 15.Ventral collar: no enlarged transverse row of scales.Ventral scales: 30 to 31 transverse rows.Sub-caudal scale rows: in uniform uninterrupted* rings.Flanks: The scales on the sides of the body are square to oval, uniform in size, andarranged in uniform longitudinal rows ( Fig 44a).




DENTITION: The teeth are long and needle-like, often extending above and below the jaws along most of the anterior jaw line. The first and fourth mandibular teeth exposedin external notches on the sides of the upper jaw. Mandibular symphysis extends to thelevel of tooth 23 to 24. The teeth are individually socketed. Formula: 5 + 23-24.

25-26

SIMILAR SPECIES: Tomistoma schlegelii , mandibular symphysis extends to the level ofthe fourteenth to fifteenth tooth.

DISTRIBUTION. Northern India, Myanmar, Nepal, West Pakistan to Assam. The riversIndus, Ganges, Mahanadi, Kaladan, and the Brahmaputra. A single record exists fromthe Irrawaddy River in Myanmar

COMMENTS. One of the most primitive, timid, and inoffensive of crocodilians, Indiangharial life figure in De Sola, 1933 is an example of Tomistoma schlegelii

REFERENCES: Boulenger, 1889; Densmore, 1983; Densmore and Owen, 1989; DeSola, 1933; Gatesy and Amato, 1992; Gmelin, 1789; Mook, 1921b; Schmidt, 1944;Wermuth, 1953; Wermuth and Mertens, 1961; Werner,1933.

Tomistom a schlegel i i (S. Muller) Malayan False GharialCONTENT: No subspecies are recognized.

SIZE: Maximum size to 500 cm. Average adult 350 cm to 400 cm. Hatchlingsapproximately 30 cm.

COLORATION: Dark reddish-brown above with dark brown or black spots and cross-bands on the back and tail. Ventrals grayish-white, with some lateral dark mottling .Juveniles are mottled with black on the sides of the jaws, body, and tail.

CONFORMATION: Snout extremely long and slender, parallel sided, length 3 to 3.5times the width at the base. Snout smooth and unornamented. The dorsal scales arebroad at mid-body and extend onto the sides of the body. The fingers are webbed at thebase. ISO’s are present on the head and body scalation.

SCALATION:Post-occipitals: frequently a single pair of slightly enlarged scales. Some individual beara number of adjoining small keeled scales; scalation divided medially by soft granularskin.Nuchals: 3 transverse rows of 2 enlarged scales, continuous with the dorsal scales.Dorsals: 22 transverse rows of 6 to 8 scales. (Note: dorsal + nuchal rows equals a totalof 22 to 23 rows).Double crest caudal whorls: 18.




Single crest caudal whorls: 17.Ventral collar: a single transverse row of enlarged scales.Ventral scales: 22 to 24 transverse rows.Sub-caudal scale rows: uniform uninterrupted rings.Flanks: One or two longitudinal rows of six to eight very enlarged scales on each side of

the body (Fig. 45).

Fig. 45

DENTITION: The first and fourth mandibular teeth are exposed in external notches onthe sides of the upper jaw. All teeth long and needle-like, interlocking on the insides ofthe jaws and are individually socketed. Mandibular symphysis extends to the level oftooth 14 to 15. Formula: 4-6 + 15-16 usually 20-21.

19-20 19 -20

SIMILAR SPECIES: Gavialis gangeticus: mandibular symphysis extends to the level oftooth 23 to 24. Crocodylus cataphractus; Crocodylus intermedius; Crocodylus johnsoni: mandibular symphysis extends to the level of the sixth to eighth tooth.

DISTRIBUTION: Indonesia, Malaysia, possibly the Viet Nam, Cambodia peninsular

COMMENTS: Biochemical genetics studies continue to place this genus within theFamily Gavialidae and not in the Family Crocodylidae.

REFERENCES: Boulenger, 1889; Densmore, 1983; Densmore and Owen, 1989;

DeRooij, 1915;DeSola, 1933; Gatesy and Amato, 1992; Loveridge, 1945; Mook, 1921b;

Muller, 1838; Schmidt, 1944; Taylor, 1970; Wermuth, 1953; Wermuth and Mertens,1961; Werner, 1933.




ACKNOWLEDGMENTS

This work is dedicated to the late Mrs. Katherine Yolla, a feisty lady who knew nothingabout crocodiles, but took pity on her son-in-law and hand typed, over and over again,my first major manuscript, “The Identification of Living Crocodilians.”

I would like to give very special thanks to my friend and colleague, Bruce Foster,Collection Manager of The Central Park Wildlife Center, WCS, who for years worked atmy side at the Bronx Zoo, collected data in Costa Rica, captured animals for me, andsaved my hide from being eaten on more than one occasion. I thank Dr. Herndon G.Dowling, and the staff of the Reptile Department: Itzchak Gilboa, Bob Brandner, andJoel Dobbin, who were always ready to help catch a crocodile for me to examine.Thanks is also due to my Brazilian colleagues, George Rebelo, INPA, and CarlosYamashita, IBAMA. To Dr. Robert Madden, Marymount College, New York, whocalculated the Caiman morphological statistics; Dr. George Amato, The WildlifeConservation Society, who conducted molecular genetics studies; Bill Meng, WCS

photographer (ret.)

There are many others too numerous to mention, who helped me in many ways overthe past 44 years. Two Crocodilian Advisory Group colleagues deserve specialpersonal thanks for their friendship and outspoken support, at times when I have takenthe unpopular road: Bill McMahan, Louisville Zoo, and Michael Davenport, National Zoo.

I also wish to thank The Wildlife Conservation Society, and especially John Behler fortheir years of support. The many museums and institutions throughout the US andBrazil, who provided specimens. The American Museum of Natural History and the lateCharles M. Bogart, Richard Zwiefel; Chicago Field Museum of Natural History; FloridaMuseum of Natural History and Wayne King; Royal Ontario Museum, Texas A&M; TheNatural History Museum of the Smithsonian Institute, George Zug and Andy Ross; TheSt. Augustine Alligator Farm, St. Augustine, Florida, and to my dear friend andcolleague, the late Fred Medem. I only wish I could send him one more box of cheapcigars.

Thanks, as well, for their great interest in crocodilian research, and support: To TheWildlife Conservation Society, The American Zoo and Aquarium Association and theCrocodilian Advisory Group, the Explorer’s Club; and especially to the U.S. Fish andWildlife Service, division of Law Enforcement, SRA Bill Donato and SRA John Meehan,WI Paul Cerniglia, Robin Lepore Esq. United States Dept. of the Interior, Office of theSolicitor; the Office of the CITES Secretariat; and the St. Augustine Alligator Farm.

Lastly to Myrna, Winnie, and the kids, who put up with my lunacy.




REFERENCES

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