/©emotrs of tbe /Duseum ot Comparative ZocJIogs AT HARVARD COLLEGE Vol LIV No ANTILLEAN TERRAPINS T Barbour and A F Carr, Jr WITH NINE PLATES CAMBRIDGE, U.S.A prtnteD for tbe /rouseum 1940 (.7^3 ANTILLEAN TERRAPINS' /•yo-7 ^'^^ would be It say that the status of the various West Indian strong possibility that this We completely satisfactory terrapins The the case is last Zoolofly APR 'V 1940 V usrar^ INTRODUCTION foolish to ^ ( word concerning the systematic There said in this paper is we present picture is is no by no means admit the inherent improbability of a new form being found in the Rio Jobabo and the common widespread form being found in the Rio Cauto in Cuba, but this seems to be the case, and no explanation can be offered but that one small population has suddenly, as where the degree of differentiation almost is zation between populations of terrapins but this whole matter aside, demonstrate We outline We is very it were, differentiated to The by no means is possibiUty of hybridi- to be cavaUerly passed understood and extremely little many lack material from too believe, however, that specific areas to difficult to more than sketch an inasmuch as we have been able to see more material than has ever been gathered together before, our conclusions are not improbably essentially correct so far as they go For many years the senior author has been collecting specimens and having them illustrated, until a considerable duties and the specialist who was fact that he could The enough so quaUfied, to be kind made had enough senior author has, however, figures, line to associate himself with this study three long visits to Cambridge He in this connection of a "finger in the pie" so that he can for the conclusions no wise dodge responsibility The stretch of the imagination pose as a with regard to studying Pseudemys led him to ask the junior author, consented so to and has in number were accumulated Administrative by no which have been reached drawings illustrating this paper, as well as some of the colored were done by Mr Eugene N Fischer who ing care with which he has illustrated colored illustrations are by the competent hand It is quite impossible to thank with material or information all of First is well known for many publications of Miss of this the painstak- Museum Other Jessie Sawyer the host of friends who have helped us and foremost Doctor Leonhard Stejneger gave us photographs of Shaw's type of Testudo rugosa no 990 in the collection of W Parker provided decussata in the British Museum the Royal College of Surgeons in London, while Mr H similar photographs of the type of Gray's Doctor Wilson Popenoe, of the Printed with the aid of a special gift Emys United Fruit Company, helped us to secure from Mr George R Agassiz memoir: museum of comparative zoology 382 The Honorable Norman Armour, formerly American material from Jamaica Minister at Port-au-Prince, and Mr F H Baker, formerly of the United States Department of Agriculture, helped us to secure specimens in Haiti Mr Charles H R M Grey In Cuba Cuban Sugar Club, Mr Henriquez, Comprador for several sugar Thrall, formerly Executive Officer of the of Soledad, Mr Norcott S companies in Havana and Agent there for Harvard University, Dr C G Aguayo and Mr Gaston selves or and J from friends S Villalba all secured terrapins for us, collecting them- in various parts of the Island Museum, V Malone aided several officers of the in procuring specimens both Messrs Wilton G Albury collecting in the on Cat Island and Great Inagua To all Bahamas, these, and others besides, our cordial thanks are given Our material from the Dominican Repubfic thank the authorities mens from Sanchez One States National of these Museum and Michigan at Ann Arbor have and their so doing is Museum of of the Field made is the is obviously inadequate and we Chicago for loaning us three speci- the type of a Museum new form The United of Zoology at the University of also kindly loaned us specimens from time to time appreciated The West Indian terrapins of the genus Pseudemys comprise a section of the wide-ranging scripta group, being most closely alUed to the Central American Pseudemys (1889), them scripta ornata who combined the This close relationship was recognized by Boulenger West Indian forms under the name rugosa and as a "variety" of Chrysemys scripta has pointed out that the scripta group is listed In a recent paper the junior author distributed, through a series of inter- grading forms, from North Carolina into South America Although each island in the Greater Antilles (and each of two of the Bahamas) has one or more recog- nizable forms, the differences between these stocks are not great; hkewise the characters differentiating the island forms from Honduranian ornata, which appears to be the annectant continental stock, are relatively sHght Indeed, in most of the distinguishing features there and were it may be found complete overlapping, not for a few minor characters which appear not to intergrade, it would seem necessary to reduce the insular forms to subspecific standing The characters Skull common The jaws and the cutting edges are to insular smooth and continental stocks are of the maxillae meet where an auxifiary notch of varying depth of the as follows —not cusped, and without marked serrations, is at the symphisis at a vertical angle, often present The alveolar surfaces upper jaw are relatively smooth, and their median ridges low and but slightly denticulate BARBOUR AND CARR: ANTILLEAN TERRAPINS The most conspicuous head Markings the orbito-mandibular The completely developed P s plastral pattern Found in old males is the most extensive in the genus are identical throughout the scripta group more or ornata appears to be less intermediate between the two sub- groups of West Indian terrapins defined later in the text by somewhat distinguished from one another differences, supratemporal and the lines on the top of the head are reduced or lacking ; The markings on the bridge and marginals Melanism stripes are the These two groups, subtle but apparently consistent combine the Cuban, Jamaican, and Cat Island forms on the one hand, and those on Porto Rico, Hispaniola, and Great Inagua on the considered, the most 383 closely, Cuban but decussata it differs is other All things perhaps the form which approaches ornata in the absence or great reduction of carapace and in the obsolescence of stripes on head and markings on the legs In general, the distribution of this group of turtles on the islands of the West Indies and the relationships with continental forms which they show, present a picture entirely compatible with what is known of the paleogeography of the Antillean region and with zoogeographic generaUties which the herpetological fauna as a whole indicates The group presents certain minor anomaUes of tribution, such as the dis- puzzUngly close apparent relationship between the Ja- maican and Cat Island forms, a relationship which appears to be much more intimate than that between P intergrading form, P barriers and time d d decussata and the strikingly differentiated but plana, but such disparity in reaction to the effects of of separation may quite properly be interpreted as a pecuUarity inherent in pseudemyd germ plasm, for the North American forms afford ample basis for such It in an interpretation seems to us quite logical to assume that the scripta complex originated and has radiated from a Mississippi Valley bordering the Embayment center, perhaps in the lowlands during Tertiary time, and that during the time of its connection with the Central American land mass, AntiUea became populated with the stock which subsequent submergence has reduced to a populations The history of the Antillean forms some conviction, but that felis) of the may in of series of insular way be inferred with two known Bahamian species (malonei and must await a more thorough understanding raphy this AntiUea and the Bahamian plateau of the relative paleophysiog- memoir: museum of comparative zoology 384 Taxonomic Arrangement In the present paper the following forms are recognized: The terrapen sub- group —Jamaica —Cat Island, Bahamas P decussata decussata (Gray) —Cuba, except Rio Jobabo and southern Pinar P terrapen (Lac^pSde) P felis Barbour del Rio : Isle of Pines P decussata plana new subspecies —Rio Jobabo, Oriente Province, Cuba —Caribbean drainage in Pinar del Rio P decussata angusta new subspecies Province, Cuba The stejnegeri sub-group P stejnegeri stejnegeri Schmidt —Porto Rico and (fide Grant 1932) Vieques Island —Hispaniola —Haiti P malonei Barbour and Carr— Great Inagua, Bahamas P stejnegeri vidua new subspecies P decorata new species In deciding between binominals and trinominals for the several forms the overlapping or failure to overlap of characters has been our only criterion Thus, the trinominal has been used for vicina which is markedly distinct from decorata, the other Hispaniolan form, but which intergrades apparently connect with stejnegeri, slightly and on and plana, for angusta since variations in decussata encroach their distinguishing characters Of the we regard scripta group, generalized, and as probably Mississippi Valley troostii as the scripta appears to be the most speciaUzed form Indian terrapins, as mentioned above, P features in common is with P most closest to the ancestral stock, while typical P s d Among s the West decussata possesses the most in the complex ornata (and also with troostii) and so may be re- garded as the most primitive of the insular forms P malonei, having departed farthest from what seems to us to be the prototypic condition, is apparently the most advanced stock We regard E vermiculata E jamao Vilar6 (Dumeril, Gray as based on the melanistic male of decussata, nomen nudum) as the non-melanistic and E gnatho Vilar6 (Cope, nomen nudum) Since Shaw's description name rugosa is The name Lac^pede is male of decussata, as the old female of decussata inadequate and his types not identifiable, the not used T palustris Gmelin is apparently a pure synonym of T terrapen barbour and carr: antillean terrapins Taxonomic Characters The characters employed of turtles are, in many by the older writers much time and paper even family characters as The West Indian Pseudemys in instances, obviously trivial clear-cut, tangible features individual used and in attempting to give diagnoses and valueless For want of and sexual variations have been repeatedly has been expended in the hsting of generic and specific diagnoses characters which we have found to be of and which have been used extensively nificance 385 more or less taxonomic sig- in the present paper, are as follows The many unsuccessful attempts Shell proportions Pseudemys on the basis of shell proportions compare comparable stages and failure to greatest help when to distinguish forms of have been due in large measure to Such sexes ratios are often of the individuals of one sex and of similar size are compared; they however, almost useless in juvenile specimens are, Stripes on head and number and nature limbs of stripes Of great value in other groups in the genus, the on the head and legs are less revealing in Indian forms, because of their exasperating tendency to lose Conformation and modelling of all Although there are no the skull West markings with age significant osteological differences in the structure of the skulls of the forms, there are slight but constant differences in the shape and proportions of the snout and in the modeUing of the surface of the we have that maxiUa It is partly distinguished the terrapen pointed out that previous writers and on the basis of such differences stejnegeri sub-groups who have used snout It should be characters have in nearly every instance been confused by the normal sexual disparity in snout length Plastral pattern In Pseudemys generally, and particularly in the scripta group, the dusky figure on the plastron, although neglected is undoubtedly of value in by most taxonomists, determining relationships The extensive rassenkreis extending from scripta in the southeastern United States to the neotropical ornata involves a progressive increase in the complexity of this pattern Inter- mediates have demonstrated the nature of the change from the pair of simple gular smudges of typical scripta to the extensive and complex figure of ornata This intricate design, as foimd in mens of Cuban decussata, and much the same in P d'orbignyi, detail in ornata, which we beUeve prove to be the southernmost extension of the scripta complex, genus, but Chrysemys is closely approached by the plastral pattern is young speci- will eventually unique for the of certain forms of memoir: museum of comparative zoology 386 Sexual Dimorphism The taxonomic confusion which —The for "rugosa" Phase more than a hundred years has sur- rounded the Antillean terrapins has been due in large part to the marked sexual dichromatism which the group displays, and to the fact that does not distinguish the sexes at The first who apphed a proper binominal name author was Lacepede (Testudo Jamaican form in 1788 The next year Shaw proposed of the the name Royal College dichromatism to one of the group terrapen, see discussion, page 393), name Lacepede's work, introduced the this all ages Gmehn T palustris for the T rugosa for a pair of specimens of Surgeons, London who described the (1789), apparently (no 990) same now unaware turtle of In 1802 in the collection These specimens, photo- graphs of which have been furnished us through the courtesy of Doctor Stejneger, what undoubtedly are the shells of One are without locaUty data is unidentifiable ; are West Indian Pseudemys, although they of the shells, apparently that of a mature female, the other, that of a male showing advanced melanism and hke- wise specifically unidentifiable, is evidently the specimen on which Shaw's figure was based and which served to estabUsh the name rugosa for melanistic males of the group no matter on which island they were found by subsequent writers Given prestige by the writings of Gray (1831, 1844, 1855, 1870, 1873) described decussata without locality and retained the name who rugosa for melanistic males, Dumeril and Bibron (1835), Sagra (1843), Gosse (1851), Strauch (1865), and others and by the plates Sowerby and Lear of (1865), the error has persisted until the present time, although the actual status of the melanistic individuals has been suggested numerous times Apparently the first writer who expressed any doubt of the distinctness of the melanistic form was Vilar6 (April and May 1867, p 119-121) who advanced the opinion that rugosa and decussata, in Cuba, were probably male and female of the same species However, help to clarify the situation : [it his subsequent remarks, quoted below, hardly should be understood that three vernacular names have been applied to the Cuban terrapins, jicotea to the females, jarico to the melanistic males, and Gundlach (1880) ambos sexos decussata, jamao apparently states, to to males which igual al de la Jicotea"] "Al hacer y otro de show no melanism or "unos individuos que tienen unas largas y la E77iys una especie de las el color Emys as de rugosa y jamao,' descansamos principalmente en un dato ' In explanation of his use of this name Vilard, in November 1867, writes as follows: "En la pagina 120 de esta tomo cito el Jamao el nombre de Emys jamao, secundum Poey, como inscrito de esta manera BARBOUR AND CARR: ANTILLEAN TERRAPINS negative, cual es el no haber hallado hasta ahora colores del Jarico Si se de la ultima." un ejemplar feminino Uega a encontrar, podrd, resultar que rugosa y decussata, sean dos buenas especies y que A few pages further on in the expresses the opinion that "el macho se 387 el las Jamao venga a same publication llama Jarico, (p los dos Jicoteas, ser el macho 104) Gundlach hembra, Jicotea." la In October of the same year Vilar6 asserts that Dumeril now agrees with him that rugosa and same decussata are the species, and introduces and describes a fourth form, E gnatho {Trachemys gnatho Cope, name was apphed This to an extraordinary phase occurring in certain females and occasionally seen decussata This in other forms manifestation of extreme age, the most striking feature augmented by a marked increase in the We of a turtle the enormous and disgreatly shortened, and width of the angle at which the maxillae meet, the whole region anterior to the eyes head is of apparently merely a is The snout becomes proportionate growth of the head to resemble the briefly nomen nudum) is relatively so have specimens broadened as hardly of decussata from several locahties exhibiting this freakish condition In two pubUcations (1875 and 1880) Gundlach makes further mention of the several forms but relegates discusses is melanism in them all to the synonymy of rugosa Fowler (1918) the Porto Rican form, and Danforth (1925) says: "There a popular idea that there are two species, a green one and a black one, but have found specimens forming all sorts of intergrades bour and Ramsden (1919) say that, "In spite species of Jicotea found in More recently De matter once and for may now popular belief, there is but one Cuba." Sola and Greenhall (1932) have undertaken to settle the all "Two in a paper entitled, The Antillean Terrapin, Pseudemys decussata.'^ of I between the two" Bar- rugosa, Species of Terrapin in Cuba; and the Cuban Terrapin, Pseudemys In the preface to this paper the reader learns that "Some solution be given to the long standing and perplexing problem concerning the terrapins of Cuba, that this remark and the multiplicity may engender teresting that all our material in the Province of is was of West Indian forms", but any optimism soon dispelled by our reading that, "It is in- collected in a restricted locality near Marti, Camaguey, Cuba, for our specimens separate into two distinct groups (!) " en los catdlogos del Sr Aug Dumeril Con mas exactitud puedo decir ahora en vista de una carta reciente Dumeril &, D F Poey, que aquel nombre aparece en una lista de los Reptiles que vivieron en el Jardin de Plantas de Paris, intercalada en una 2" noticia publicado por Dumeril en los Archives del Museo, t X, en Setiembre de 1861, p 435, la inscrlpcion de num 27, Ermjs Jamao, A Dum., segun del Sr F Poey, macho y hembra, Habana, individuos." memoir: museum of comparative zoology 388 One of these "distinct groups", the old name P The rugosa 44876) described under the are merely those it "an adult male" as are the other specimens which they men- is and tion as having examined, plate of rugosa which De finally, reference is made to Sowerby and Lear's Sola and Greenhall reproduce and which some indeterminate melanistic phase of is is specimen selected as typical of the form (American of a melanistic male, the Museum male group, characters presented as distinguishing species of merely the is West Indian Pseudemys, and also obviously a male P decussata known is summarily dismissed with the remark that range its is "only Rio Tana and Rio Jobabo in the Province of Camaguey, Cuba," and as that "all the specimens in this group agree with the original description of that adequately differentiates P decussata from P rugosa," followed tion of these characters which young males from melanistic Gray by a quota- really nothing but distinguish females We individuals are then referred to and Sowerby and Lear's plate of decussata, also reproduced, which probably really does represent the Cuban form It is a female Next there comes a "Key to the Cuban Species of a satisfactory key to the sexes of nearly any of the finally Pseudemys" which West Indian we read that "Some confusion has been caused by a many to recognize naturaUsts, offered by way is who beUeve of proof a in the presence of failure and on the part of two species on male and a female of the same also encountered in our not be amiss." quite the sex characters present in Pseudemys For example, several Cuban of error is terrapins, The time-honored own tail species their island, This type country, so the following key and may toe-nail sex characters are then presented In 1934 Grant and De Sola approach the problem from a shghtly different angle, the argument being that there are two species of decussata, approximately throughout the Antilles coextensively distributed But here again rugosa is Pseudemys, rugosa and nothing more than the old male really nothing at all surprising about the Greater —so there two species having coextensive is dis- tribution Grant and De Sola point out that De Sola and Greenhall state that "the nails of the fore feet of after males of decussata are elongated", but that "their figure Sowerby and Lear does not show the nails in the figure are not long enough nails long for a male, enough." but this is the delineator or the Uthographer, since the specimen figured a female It is true that the no indictment of is quite obviously PLATE Plate Miss J Sawyer del Dorsal and ventral view of Pseudemys decussata decussata, Rio Cauto, Cuba, Mus Comp Zool No 34136 Melanistic cf MEM MUS COMP ZOOL BARBOUR AND CARR ANTILLEAN TERRAPINS PLATE PLATE Plate Read of E N Fisher del Pseudemys felis, C&t Island Mus Comp Zool No 38385 Ad Head of Pseudemys Young Head of Head of Head Pseudemys terra-pen, decorata, Pseudemys near Kingston, Jamaica Fond Parisien, Haiti stejnegeri stejnegeri, No 37295 Young Mus Comp Zool Rio Piedras, Puerto Rico Zool No 39704 No 36863 Ad Mus Comp cf Zool cf Pseudemys decussata decussata, melanistic, old Zool No 34136 of Mus Comp cT, Rio Cauto, Cuba Mus Comp MEM MUS COMP ZOOL BARBOUR AND CARR ANTILLEAN TERRAPINS PLATE "S^.yT Q^- r/^^ r PLATE E N Fisher del Plate Head Pseudemys decussata plana Rio Jobabo, Cuba Mus Comp Zool No 34133 of Head of Pseudemys Young Head of Ad decussata decussata Soledad, Cuba Mus Comp Zool No 34147 Pseudemys decussata plana Rio Jobabo, Cuba Mus Comp Zool No 34128 cf Foot of Pseudemys decussata decussata, Rio Cauto, Cuba Mus Comp Zool No 34136 Mus Comp Zool No 34136 Melanistic cf Head of Pseudemys decussata decussata, Rio Cauto, Cuba Melanistic cf Foot of Pseudemys decussata decussata, Soledad, Cuba Mus Comp Zool No 34147 Young C&uto, Cuba Mus Comp Zool No 34137 Rio Jobabo, Cuba Mus Comp Zool No 34132 Head oi Pseudemys decussata decussata, Jiio Melanistic cf Foot of Pseudemys decussata plana Young MEM MUS COMP ZOfiL 3ARBOUR AND CARR ANTILLEAN TERRAPINS PLATE ^' ' PLATE Plate Under surface Zool Under del Pseudemys decussata decussata Soledad, Santa Clara, Cuba Mus Comp No 34177 Young Upper view Zool of E N Fisher of Pseudemys decussata decussata Soledad, Santa Clara, Cuba Mus Comp No 34177 Young surface of Pseudemys decorata Thomazeau, Haiti Mus Comp Zool No 36855 Zool No 36855 Young Upper view Young of Pseudemys decorata Thomazeau, Haiti Mus Comp MEM MUS COMP 200L BARBOUR AND CARR ANTILLEAN TERRAPINS PLATE ... possibility that this We completely satisfactory terrapins The the case is last Zoolofly APR 'V 1940 V usrar^ INTRODUCTION foolish to ^ ( word concerning the systematic There said in this paper... United States National locality Museum San Juan, Porto Rico 25642, mature female {,7^3 ^Pfi 24 1940 ERRATA Page 407 Plastrons of Pseudemys d Caption of Fig should read: stejnegeri and s young