Three New of venusta (Testudines: ) from Honduras, Northern Yucatán (Mexico), and Pacific Coastal Panama

By William P. McCord1, Mehdi Joseph-Ouni2, Cris Hagen3, and Torsten Blanck4 1East Fishkill Hospital, Hopewell Junction, NY, USA; 2EO Wildlife & Wilderness Conservation, Brooklyn NY, USA; 3Savannah River Ecology Laboratory, Aiken, SC, USA; 4Forstgartenstr 44, Deutschlandsberg, Austria

Abstract. Upon examination of live and preserved specimens from across the range, several unnamed distinct forms of Trachemys venusta (Gray, 1855) were recognized, leading to the description here of three biogeographically isolated, morphologically distinct subspecies: Trachemys venusta uhrigi ssp. nov., Trachemys venusta iversoni ssp. nov., and Trachemys venusta panamensis ssp. nov. Head and neck stripes, along with cara- pacial and plastral patterns are critical to identification in this group. Formal descriptions and diagnoses are given herein. Other Central American Trachemys are also discussed for comparison.

Keywords: , emydid, Trachemys venusta uhrigi ssp. nov., Trachemys venusta iversoni ssp. nov., Trachemys venusta panamensis ssp. nov., Honduras, Mexico, Panama, Meso-America.

Slider of the Trachemys GRAY (1855) declared eight syntypes for his “Venus- Agassiz, 1857, range from north and east of like” without designating a holotype. However, the Rio Grande in the United States, in 1873 he referred to only one syntype as “Emys through Mexico, Central America, the venusta”: stuffed specimen “e” (1845.8.5.26) in the West Indies, and South America as far as British Museum of Natural History, labeled “Charming northeastern Argentina. IVERSON (1985) resurrect- Emys” for its beautiful pattern — this reference led ed Trachemys from synonymy with (Gray, SMITH and SMITH (1979) to designate BMNH 1855). SEIDEL (2002) elevated Trachemys scripta 1845.8.5.26 (Dyson collection) as the species lectotype, venusta (Gray, 1855) to full specific status with three retaining Gray’s original uncertain type locality subspecies (see below). For taxonomic changes “Honduras” GRAY (1855). Specimen “e” (1845.8.5.26) between original descriptions and present usage of all also appears to be illustrated in “TAB XII.A.” species and subspecies in this paper, see the synonomy (GRAY, 1855), further supporting its designation lists in FRITZ and HAVAS (2007). Dispersal of (by Code recommendation) as lectotype. North American ancestral stock along coastal regions Six of the eight Trachemys venusta syntypes and across the interior in narrow or lowland areas has (1839.12.26.70, 1844.2.19.3, 1845.10.25.17, 1849.12.7.4, led to widespread distribution of T. venusta with mul- 1849.7.28.25, and 30G) have Belize-like plastral pat- tiple subspecies on both coasts of Central America terns and/or head stripes (see below); one other (30B) (LEGLER, 1990). is morphologically too vague to identify. [Note: Further discussion of Trachemys venusta requires an specimen “f” (Gray, 1873) BMNH 1845.10.25.19 is an understanding of exactly what Gray described in 1855. iguana; Gray intended to give no. 1845.10.25.17 to a

Symphyseal Stripe Postorbital Stripe Secondary Mandibular- Cervical Stripe Central Secondary Orbitocervical Stripe Mandibular Stripe Primary Orbitocervical Stripe Paramedian Ventral Neck Stripe

Median Ventral Neck Stripe

Head and neck stripes: note Y figure on chin, and that the mandibular stripe joins the secondary mandibular-cervical stripe. Photos: H.-D. Philippen

REPTILIA 39 supposed “Honduran” Dyson turtle venusta grayi’s range) for the follow- specimen (Colin McCarthy, pers. ing reasons (quotes are from the comm.) that we feel has a Belize- original description): shell “nearly like plastral pattern]. The lectotype two feet long” (T. v. grayi reaches 60 (1845.8.5.26) has lost its plastral and cm; Caribbean Hondurans reach 30 bridge patterns, but the ventral cm; Uhrig collection; SMITH and neck stripes and position and quali- SMITH, 1979); “jaws serrate in the ty of the costal ocelli also suggest a same manner as the E. mobilensis” Belize origin. (PRITCHARD, 1979, states “with SCHMIDT (1941) stated that all Pacific Pseudemys, the jaw “all of the Dyson collection in the edges are markedly serrate” — the British Museum, labeled only jaws of Atlantic or Caribbean spec- “Hondurus,” should be referred to imens are smooth or finely ser- British Honduras” (now Belize). rate); and the median and parame- Apparently Dyson (1845) snake dian ventral neck stripes are of specimens from “Honduras” equal width, as is also the case with proved to be of Belize origin. T. v. grayi (paramedian stripes are According to Dr. Colin McCarthy wider than median stripes in (BMNH), the lectotype (1845.8.5.26) Caribbean Hondurans). Consid- was part of a series (1845.8.5.14–39) ering the T. v. grayi-like diagnostic “from Belize,” that was “pur- features of Emys valida, the Trachemys ornata. Photos: D. MacKinnon chased from Cuming, collected by emended locality of Emys valida Dyson.” The designation “Belize” to Pacific coastal Honduras, and was not dittoed for the locality of mention by PRITCHARD (1979) every specimen (even of same that he was given a T. v. grayi cara- species) in the original registry, pace in Tegucigalpa, Honduras, probably for sake of expediency, we herein designate Emys valida even though the source “Dyson” is LeConte, 1859 a of T. v. dittoed for all. This led an earlier grayi. Furthermore, according to curator to list the entire series SMITH and SMITH (1979), “valida 1845.8.5.14–39 as from Belize in his was never adopted as valid after notebook for the museum. its description in 1859.” There- We conclude that GRAY (1855) fore, we herein designate Emys was mistaken in giving “Hon- (Ptychemys) valida LeConte, 1859, Trachemys ornata: note widening of postorbital duras” as locality for specimen “e” as a nomen oblitum to avoid taxo- stripe; the mandibular stripe joins the primary orbitocervical. Photo: R. Gurley (1845.8.5.26), now the lectotype. nomic instability. Therefore, we herein emend the Thirty years after Gray described Sinaloa, southward to the region of type locality of Emys venusta, Emys venusta, GÜNTHER (1885) Acapulco, central Guerrero. In Gray, 1855, to Belize (= British described Emys salvini with type Trachemys ornata the postorbital Honduras), thus allowing taxo- locality “Guatemala.” The holo- stripe starts thin at the orbit and nomic continuity and stability, type is BMNH 1946.1.22.76 (for- then expands (widens) over the while satisfying Gray’s original merly 64.2.19.5) and is depicted in tympanum, making it the dominant intention. “Tabb. II & III.” as drawings in the head stripe; the mandibular stripe Next we resolve the status of two book with the description. We find joins the primary orbitocervical synonyms of Trachemys venusta that this specimen morphologically stripe (or sometimes a secondary (Gray, 1885). Four years after matches the Belize T. venusta mandibular-cervical stripe); the Gray described Emys venusta, (Gray, 1855), making E. salvini symphyseal stripe joins the parame- LECONTE (1859) described Emys Günther, 1885, a subjective junior dian neck stripes forming a Y- valida with the type locality synonym of T. venusta (Gray, shaped marking on the chin; the “Honduras.” The holotype (Acad. 1855). mandibular tomium is coarsely ser- Nat. Sci. Philadelphia 216) consists Until MOLL and LEGLER (1971) rate; the carapace reaches 38 cm, of a head, two front feet, and a tail. recognized Pseudemys scripta venusta and is widest at the seam between The shell was destroyed in transit (= Trachemys venusta today), most the 7th and 8th marginal scutes; and is not available. We herein authors considered it a synonym of costal ocelli occupy the posterome- emend the type locality of Emys Trachemys ornata (Gray, 1831). dial surface of the scute, are dark valida LeConte, 1859, to Pacific Trachemys ornata sensu stricto centered, irregular, often incom- coastal drainages of Honduras (the occurs only along the Pacific coast plete, and of a diameter often half southeasternmost end of Trachemys of Mexico from Culiacán, northern or less than half the width of the

REPTILIA 40 Belize (= British Honduras). The species ranges through Central America from northern Tamaulipas, Mexico on the Gulf coast, eastern Oaxaca, Mexico on the Pacific coast, to southern Panama on both its Caribbean and Pacific coasts. SEI- DEL (2002) designated three sub- species for T. venusta: T. v. venusta, T. v. grayi (Bocourt, 1868), and T. v. Trachemys emolli: note fading of secondary carapace scute patterns; postorbital stripe does not contact orbit cataspila (Günther, 1885) — these and is constricted on tympanum. On the plastron note the “bow-tie” figure at the anterior. Photos: K. Edwards subspecies had all previously been members of the Pseudemys scripta subspecies complex, belonging to the ornata group. The subspecies T. v. venusta sensu stricto ranges (restricted herein) from just south of Punta del Morro, southward through Vera- cruz, Oaxaca, and Tabasco, through the southern Yucatán Peninsula (Campeche and Quintana Roo), Trachemys venusta venusta: note widening of costal ocelli laterally; costal ocelli almost fill the scutes. Mexico, northern Guatemala, On the plastron note central fading, leaving an empty double-lined figure. Photos: D. Uhrig Belize, and into Caribbean coastal scute; the first vertebral scute is the median neck stripe; primary Guatemala. In this taxon the pos- wider than long; there are two or costal ocelli occupy 25–50% of the torbital stripe is yellow, moderately three light stripes on each bridge; scute on its posteromedial surface; thin, and contacts the orbit; the pri- the adult plastral pattern consists of secondary costal ocelli fade in mary orbitocervical stripe is two to a medial pair of dark lines on each adults so that only the often incom- three times as wide as the postor- scute surrounding an open area plete primary costal ocelli remain bital stripe and is the dominant with lateral extension on 50% of surrounding dark centers; sec- head stripe; the mandibular stripe is the remaining seam extremities; ondary orange markings on the isolated; the symphyseal stripe does hatchlings may also have a median vertebral scutes also fade with age; not connect with the paramedian seam-following pattern, the overall the carapace reaches 40 cm, and is neck stripes, so no Y is present on pattern being squared-off at both widest at the 7th marginal; the first the chin; the symphyseal and medi- ends with a posterior dip in the pat- vertebral scute is longer than wide; an neck stripes are wider than the tern on the lateral humeral/pectoral the bridges have two light lines; the paramedian stripes; the iris is yel- seam; the plastral seam formula plastral pattern of hatchlings is low; the mandibular tomium is fine- starts with abd > an > fem; rump greatly reduced, with bold dark ly serrate; bright orange primary pattern consists of 25–30 yellow lines longitudinally following close- costal ocelli are very large, bolder blotches or spots aligned vertically. ly both sides of the median scute laterally, light orange centered, LEGLER (1990) described seams, but not the intergular seam, complete on the first three scutes, Trachemys emolli from Lakes and only the anterior half of the and fill each scute 95–100% trans- Managua, Nicaragua and adjacent interanal seam. Fine dark lines versely, occupying 50–90% of the waters. In this taxon, the postor- also extend to the full extent of all posterior scute surface; the cara- bital stripe does not contact the lateral plastral seams. With age the pace reaches 48 cm and is widest at orbit, and is relatively wide with a pattern changes to irregularly the 7th marginal; the first vertebral constriction over the tympanum cover the central 50–60% of the scute is wider than long; secondary that may form a bi-lobate marking plastron (lateral extensions fade), costal and vertebral scute markings or may terminate the stripe creat- with an often isolated bowlike fig- (basically parallel to the median ing an isolated oblong or round ure developing on the gular/humer- line) do not totally fade. There are orange spot; the postorbital stripe al scute seams; the plastral seam two light lines on each bridge. The is the dominant head stripe; the formula starts with abd > an > gul; plastral pattern of hatchlings is mandibular stripe is isolated; the the central stripe on the rump is greatly reduced (20% of plastral symphyseal stripe is connected to usually broken. surface) with bold dark lines longi- the paramedian neck stripes form- In 1855 Gray described Emys tudinally following closely both ing a Y on the chin; the ventral venusta (= Trachmys venusta). As sides of the median scute seams but paramedian stripes are wider than emended above the type locality is not the intergular seam, and only

REPTILIA 41 stripe connects with the paramedi- abd > an > gul. There is no central an neck stripes and forms a Y on stripe on the rump. the chin; the median and paramedi- In 1885 Günther described Emys an neck stripes are equal in width; cataspila (= T. v. cataspila), which the mandibular tomium is fine to ranges from the Rio San Fernando moderately serrate; the snout is drainage of northern Tamaulipas often protuberant in both sexes, south to the vicinity of Punta del more so in males; light yellow costal Morro, Mexico, where the Sierra ocelli are dark centered, often Madre Oriental separates it from T. incomplete, and are located in the v. venusta. In this taxon the postor- Trachemys venusta grayi: melanism has obliterated posteromedial corner of the scute bital stripe contacts the orbit, start- all scute patterns. Photo: R. Gurley surface, with a diameter about 50% ing thin and yellow to orange, and the anterior half of the interanal of the width of the scute. Melanism expanding over the posterior tym- seam. Fine dark lines also extend obliterates the entire carapacial panum to an orange patch, making on the lateral seams (all but the pattern in adults. The carapace it the dominant head stripe; it then gular/humeral), varying tremen- reaches 60 cm (OBST, 1985), is turns yellow as it thins and contin- dously from 0–90% of seam length. widest at the seam between the 7th ues down the neck. There is often a With age the plastral pattern and 8th marginals; the first verte- break in the postorbital stripe over becomes a medial pair of dark lines bral scute is longer than wide. the anterior tympanum. The prima- surrounding an open area on each There are two light lines on each ry orbitocervical stripe is twice the scute, and the lateral extensions bridge. The plastral pattern of width of the anterior postorbital fade. The plastral seam formula hatchlings is moderately reduced, stripe; the mandibular joins the pri- starts with abd > an > gul (usually) with bold dark lines longitudinally mary orbitocervical stripe; the sym- or fem. The central stripe on the following both sides of the median physeal stripe joins the paramedian rump is solid. As with all T. venusta scute seams (except the intergular stripes and forms a Y on the chin; the sexes are close in size and the and interhumeral, and only the the median and paramedian stripes foreclaws are the same (short) in anterior 75% of the interanal of the ventral neck are equal in males and females. seam). Dark swirling lines cover width; the mandibular tomium is In 1868 Bocourt described Emys only the medial 50% of the pec- finely serrate or smooth; orange to grayi (= T. v. grayi), which ranges toral/abdominal, abdominal/femora red costal ocelli are dark centered, from the region of Salina Cruz in l, and femoral/anal seams. Double often incomplete, and are located in eastern Oaxaca, southeastward bold dark lines also extend from the posterolateral corner of the through coastal Chiapas, Mexico, the mid-intergular seam to the scute surface. The diameter of the to Pacific coastal Honduras (see medial half of the gular/humeral boldest costal ocelli is ~40% of the PRITCHARD, 1979). In this taxon seam, and drop mid-seam posteri- width of the scute. In many speci- the postorbital stripe contacts the orly over the medial third of the mens there are multiple ocelli (com- orbit and is a pale yellow, consis- humeral scutes, connecting with plete or partial) with dark blotches tently thin stripe; the pale yellow bold dark lines (continued laterally on the first and second costals. primary orbitocervical stripe (often by fine dark lines) on the medial Secondary orange markings are splitting to outline the anterior, humeral/pectoral seams, forming a plentiful and chaotic. The vertebral dorsal, and ventral tympanum) is boxlike figure at the anterior of a scutes often each have an elongated 50% wider than the postorbital generally irregular pattern. The median black blotch. The black stripe making it barely the domi- entire plastral pattern fades and is blotches of the marginal scutes are nant head stripe; the mandibular subject to melanism with age. The usually on the posterior of each stripe is isolated; the symphyseal plastral seam formula starts with scute, anterior to the intermarginal

Trachemys venusta cataspila: note multiple primary ocelli on the first costal scutes; Trachemys venusta cataspila: note the break in the postorbital stripe before it primary costal ocelli located posterolaterally on the scute; and teardrop-like widens over the tympanum and that the mandibular stripe joins the primary figures on the lateral humeral/pectoral seams. Photo: D. MacKinnon orbitocervical. Photo: D. Uhrig

REPTILIA 42

seam. The carapace reaches 32 cm, is widest at the 7th marginal, and the first cervical scute is wider than long. There are two light lines on each bridge. The plastral pattern of hatchlings is a reduced or moder- ately reduced pattern of bold dark lines following all longitudinal and horizontal seams of the plastron except the anterior 80–90% of the intergular seam, sometimes the pos- terior 10–30% of the interanal seam, and sometimes the outer (lat- eral) 10% of most horizontal seams. The bold dark lines always reach the full lateral extent of the humer- al/pectoral seams where there is a notable, lateral, posteriorly orient- ed dark rounded figure. The gular/humeral seams often have a simple forked appearance, but a Trachemys venusta uhrigi ssp. nov. holotype: note consistently thin costal ocelli, greatly expanded boxlike pattern at the anterior of plastral pattern, and four light stripes on bridges. Photos: W. P. McCord the plastral figure is not uncom- mon. In adults the plastral pattern diffuses and fades, with the pattern of the humeral/pectoral seam and the gulars often remaining. The plastral seam formula starts with abd > an > fem. The central stripe on the rump is broken into multiple varying lines and yellow blotches. After many years of working with Trachemys venusta we present here original formal descriptions of three new subspecies. This is pub- lished to provide a public and per- manent scientific record. Date of publication: Reptilia (GB) no. 71 (ES no. 84, and IT no. 33), Castelldefels, Spain, mailed 02 August 2010.

UHRIG’S SLIDER TURTLE Trachemys venusta uhrigi ssp. nov.: note thin neck stripes. Photo: D. Uhrig December 1976 by Diderot F. Gicca Suspected to range along the Trachemys venusta uhrigi ssp. nov. in the Río Ulúa, 8.0 km south of remaining Caribbean coast of Testudines Linnaeus, 1758. San Pedro Sula, Honduras. Honduras, Nicaragua (LEGLER, Suborder Cope, 1868. Type locality: Río Chamelecón 1990, Fig 7.6, depicts a T. v. uhrigi in Emydidae Rafinesque, 1815. drainage 3 km south of San Pedro Caribbean coastal Nicaragua as P. s. Holotype (designated herein): Sula, northwestern Caribbean venusta), Costa Rica, Panama, and Florida Museum of Natural History coastal Honduras. into the lower drainages of the (FLMNH) 157800, adult female, Distribution: Presently known by Atrato River of northwestern preserved in alcohol, collected in these authors from the vicinities of Colombia. May 1993 by Dennis Uhrig in the San Pedro Sula and La Ceiba in Etymology: Named in honor of Río Chamelecón, 3.0 km south of northwestern Caribbean coastal Dennis Uhrig, who has dedicated San Pedro Sula, Honduras. Honduras eastward to the lower his life to studying and working Paratype (designated herein): Patuca River drainage, in the region with all Trachemys and Pseudemys FLMNH 105425, subadult female, of northeastern Caribbean coastal turtles, and who was the first to preserved in alcohol, collected in Honduras called “La Mosquitia.” notice the uniqueness of this turtle.

REPTILIA 43

Description femoral/anal seams. The plastral Plastron. There are usually four In Trachemys venusta uhrigi the pattern of older adults fades from light lines on each bridge of T. v. postorbital stripe contacts the orbit the center first, eventually with only uhrigi; two in T. v. venusta. In juve- and is yellow and consistently thin; remnants on the gulars, humerals, nile T. v. uhrigi the plastral pattern the primary orbitocervical stripe is and pectorals. The plastral seam is greatly expanded to cover ~95% 25–50% wider than the postorbital formula starts with abd > an > fem. of the plastral surface; in T. v. stripe making it barely the domi- The central stripe on the rump is venusta the pattern is greatly nant head stripe; the mandibular usually broken into three or four reduced, involving ~20% of the stripe is isolated; the symphyseal yellow segments. plastral surface. The plastral seam stripe does not connect with the formula of T. v. uhrigi starts with paramedian neck stripes, so no Y is Diagnosis abd > an > fem; usually abd > an > present on the chin; the paramedian We herein differentiate Trachemys gul in T. v. venusta. The central neck stripes are wider than the venusta uhrigi ssp. nov. from the stripe on the rump of T. v. uhrigi is median stripe; there is often a trans- nominotypical (conspecific) form T. broken; solid in T. v. venusta. verse yellow stripe connecting the v. venusta sensu stricto (as described Trachemys v. uhrigi can be differ- two paramedian stripes on the and with the range given above), and entiated from T. v. grayi as follows: upper ventral neck forming an H- its two neighboring forms T. v. grayi Head and neck. In T. v. uhrigi, like figure; the iris is bluish green; (conspecific) and T. emolli (con- the yellow postorbital stripe is unin- the mandibular tomium is smooth generic). Of the characters given in terrupted over the tympanum; in T. or finely serrate. Hatchlings have a the foregoing descriptions, only those v. grayi it is pale-yellow, and often pale yellow to orange ocellus differentiating T. v. uhrigi from T. v. splits and outlines the tympanum. (incomplete posteriorly), around a venusta, T. v. grayi, and T. emolli are In T. v. uhrigi the paramedian ven- green-centered (matches carapace) given here. tral neck stripes are usually wider small black ring posterocentrally Trachemys v. uhrigi can be differ- than the median stripe; equal widths located on each costal and vertebral entiated from T. v. venusta as follows: in T. v. grayi. The symphyseal stripe scute. In adults, the consistently Head and neck. In T. v. uhrigi the of T. v. uhrigi does not connect with thin, pale yellow to orange (com- postorbital stripe is very thin; the the paramedian stripes; with T. v. plete or incomplete posteriorly) pri- primary orbitocervical stripe is also grayi it does, forming a Y on the mary ocelli fill 25–50% (of the pos- thin (only 25–50% wider than the chin. T. v. uhrigi has a finely serrate teromedial surface) of the costal postorbital); the paramedian stripes tomium; moderately serrate in many scutes. Each primary ocellus has a are usually wider than the median T. v. grayi. In T. v. uhrigi the snout usually solid black blotch in the ventral neck stripe and the iris is is more pointed in males than center, plus two or three incom- bluish green. In T. v. venusta the females; in T. v. grayi, the snouts of plete, thin, faded secondary costal primary orbitocervical stripe is much both sexes are often pointed. The rings that fill the anterior and later- wider than (2–3 times) the moder- iris is bluish-green in T. v. uhrigi; yel- al areas of the scute not filled by the ately thin postorbital stripe; the low in T. v. grayi. primary ocellus. The vertebral median stripe is the widest on the Carapace. In T. v. uhrigi the cara- scutes in adults display thin, faded ventral neck, and the iris is yellow. pacial pattern remains for life; in T. irregular triangular figures. The Carapace. The yellow to orange v. grayi melanism obliterates all marginals bear dark areas centered costal ocelli and secondary mark- patterns with age. The carapace on the intermarginal seams. The ings are very pale on hatchling T. v. length of T. v. uhrigi is known to carapace reaches 30 cm in length, is uhrigi; bolder orange on hatchling reach 30 cm; 60 cm in T. v. grayi. widest at the seam between the 7th T. v. venusta. The black costal The carapace is widest at the 7th and 8th marginals, and the first ver- rings within the ocelli (also on mar- marginal in T. v. uhrigi; at the seam tebral scute is wider than long. ginals and vertebrals) of hatchlings between the 7th and 8th marginals There are three or four light lines are shell-green centered in T. v. in T. v. grayi. In T. v. uhrigi the first on each bridge. The plastral pat- uhrigi; orange in T. v. venusta. In vertebral scute is wider than long; tern of hatchlings is a greatly adults the primary costal ocelli of longer than wide in T. v. grayi. expanded pattern of bold dark lines T. v. uhrigi are consistently thin, Plastron. In T. v. uhrigi the plas- involving 90–95% of the total plas- pale, solid black centered, and tral pattern is greatly expanded; in tron. All but the anterior 15–20% occupy 25–50% of the posterome- T. v. grayi it is moderately reduced of the gulars, posterior 10–20% of dial surface of the scute; in T. v. with a boxlike figure at the anterior the anals, and the lateral 20% of venusta, bright orange primary aspect. The plastral seam formula the humeral, pectoral, abdominal, costal ocelli (bolder laterally), hav- of T. v. uhrigi starts with abd > an > and femoral scutes are covered ing orange within the black cen- fem; in T. v. grayi, abd > an > gul. with a swirling pattern reaching the ters, occupy 50–90% of the posteri- T. v. uhrigi has a segmented central lateral extremities of the humer- or surface of each scute, and stripe on the rump; T. v. grayi has al/pectoral, abdominal/femoral and 95–100% transversely. no central stripe.

REPTILIA 44

Trachemys v. uhrigi can be differ- seam between the 7th and 8th mar- 13.8 km east of Buctzotz, Yucatán, entiated from T. emolli as follows: ginals. The first vertebral scute of Mexico. Head and neck. In T. v. uhrigi T. v. uhrigi is wider than long; Paratype (designated herein): the consistently thin postorbital longer than wide in T. emolli. FLMNH 50476, subadult female, stripe connects with the orbit; in T. Plastron. T. v. uhrigi has three or preserved in alcohol, collected in emolli it does not reach the orbit, four light lines on each bridge; T. April 1981 by John B. Iverson in and as the stripe widens over the emolli has two. The plastral pat- Cenote Xkolar, 16.6 km northeast tympanum there is a constriction tern of T. v. uhrigi is greatly of Izamal (toward Tunkas), creating a bi-lobed appearance, or expanded; in T. emolli it is greatly Yucatán, Mexico. sometimes terminating the stripe reduced, with a bow-like figure at Type locality: A Cenote on the after the first lobe. The primary the anterior aspect in adults. In T. north side of the highway, 13.8 km orbitocervical stripe is dominant in v. uhrigi the plastral seam formula east of Buctzotz, Yucatán, Mexico. T. v. uhrigi; postorbital in T. emolli. begins with abd > an > fem; in T. Distribution: Presently known In T. v. uhrigi the symphyseal stripe emolli, abd > an > gul. from the Cenotes in northern does not connect with the paramedi- Yucatán State, Mexico. an neck stripes; in T. emolli it does, IVERSON’S SLIDER TURTLE Etymology: Named in honor of forming a Y on the chin. Taxonomy John B. Iverson, a Chelonian spe- Carapace. In T. v. uhrigi all sec- Trachemys venusta iversoni ssp. nov. cialist of the highest caliber, who ondary scute markings remain in Order Testudines Linnaeus, 1758. did much field work with this group adults; in T. emolli they fade, leav- Suborder Cryptodira Cope, 1868. of turtles. ing only primary markings that are Family Emydidae Rafinesque, 1815. more brilliantly colored and wider Holotype (designated herein): Description than in T. v. uhrigi. T. v. uhrigi FLMNH 50478, adult female, pre- In Trachemys venusta iversoni reaches ~30 cm in length; T. emolli, served in alcohol, collected in April the postorbital stripe contacts the 40 cm. T. v. uhrigi is widest at the 1981 by John Iverson in a Cenote orbit as a very thin stripe, then 7th marginal; in T. emolli, at the on the north side of the highway, expands to a dull orange patch

Trachemys venusta iversoni ssp. nov. holotype: note greatly expanded plastral pattern, and two light stripes on bridge. Photos: G. Salmon

Trachemys venusta iversoni ssp. nov.: note posterocentral location and consistent width of the primary costal ocelli; golden-yellow iris; bluntness of face; boldness of secondary stripes; and widening of postorbital stripe over tympanum. Photos: J. B. Iverson

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over the tympanum, then narrows reaches the posterior end of the is known to reach 20 cm, and is again and continues down the neck median plastral seam in both widest at the seam between the as a moderate stripe; the primary hatchlings and adults. The plastral 7th and 8th marginals; 48 cm for T. orbitocervical stripe is two to three seam formula varies: 65% of speci- v. venusta, widest at the 7th mar- times the width of the postorbital mens starting with abd > an > pect; ginal. The secondary costal and stripe making it the dominant head 35% starting with abd > an > fem. vertebral scute markings of T. v. stripe; secondary facial stripes are The central stripe on the rump is iversoni fade with age; not in T. v. prominent; the mandibular stripe is usually broken. venusta. isolated; the symphyseal stripe Plastron. The plastral pattern of does not connect with the parame- Diagnosis T. v. iversoni is greatly expanded; dian neck stripes, so no Y is present We herein differentiate T. v. greatly reduced in T. v. venusta. on the chin; the symphyseal and iversoni ssp. nov. from the nomino- The plastral seam formula of T. v. median ventral neck stripes are typical form T. v. venusta sensu iversoni begins with abd > an > pect wider than the paramedian stripes; stricto (as described herein and in 65% and abd > an > fem in 35% the iris is golden-yellow; the with the range given above). Of the of specimens studied; in T. v. venusta, mandibular tomium is finely ser- characters given in the foregoing abd > an > fem in 80%, and abd > rate; the snout is blunt in both description, only those differentiat- an > gul in 20%. The central stripe sexes; in hatchlings, pale yellow- ing T. v. iversoni from T. v. venusta on the rump of T. v. iversoni is usu- orange ocelli surrounding solid are given here. ally broken; always solid in T. v. black spots (light centered in Head and neck. The postorbital venusta. some) are located posterocentrally stripe of T. v. iversoni widens over The plastral pattern of T. v. on each costal scute surface; adults the tympanum; consistently thin in iversoni can be differentiated from have consistently moderately wide T. v. venusta. The coloring of all the also very expanded pattern of T. pale orange costal ocelli filling the head stripes are a pale yellow- v. uhrigi as follows. In T. v. iversoni posterocentromedial 25–30% of orange in T. v. iversoni; bold yellow the plastral pattern reaches the pos- the costal scute surfaces; primary in T. v. venusta. The secondary terior end of the median seam in costal ocelli do not contact the ver- head stripes are more prominent in both hatchlings and adults; in T. v. tebral or marginal scute seams; sec- T. v. iversoni than T. v. venusta. uhrigi it reaches the posterior end ondary ocelli fill the anterior and The iris is golden-yellow in T. v. in hatchlings only. In T. v. iversoni lateral costal scutes; primary verte- iversoni; yellow in T. v. venusta. juveniles, the lateral pattern on the bral scute markings are often sym- The anterior face of T. v. iversoni is humeral/pectoral seams is oriented metrically aligned (parallel) with blunter (in both sexes) than in T. v. posteriorly and ends on the pectoral the median stripe; marginals bear venusta. scute; in T. v. uhrigi the humeral/ dark areas centered on the inter- Carapace. The pale orange, rela- pectoral seam pattern is oriented marginal seams; carapace length is tively small costal ocelli of T. v. anteriorly and ends on the humeral known to reach 20 cm, and is iversoni occupy 25–30% (in the scute. widest at the seam between the 7th posterocentromedial surface) of Apparent intergradation, pre- and 8th marginals; the first verte- the overall scute; in T. v. venusta senting mixed characters of both T. bral scute is usually longer than the large bold orange costal ocelli v. iversoni and T. v. venusta was wide (sometimes equal). There are occupy 50–90% (of the entire pos- found as far north as Muna in the two light stripes on each bridge. terior surface) of the overall scute northwestern Yucatán Peninsula The plastral pattern of hatchlings is surface. In T. v. iversoni the stripe and Cobá in the northeastern a greatly expanded pattern of bold width of the primary costal ocelli Yucatán Peninsula, both at ~20.5 dark lines involving 90% of the is consistent throughout; in T. v. degrees north latitude. total plastron. All but the anterior venusta it is wider laterally. The 20–25% of the gulars, the posterior primary costal ocelli of T. v. PANAMANIAN SLIDER TURTLE 20–25% of the anals, and the later- iversoni never contact the vertebral Taxonomy al 20–25% of the humerals, pec- or marginal seams; in T. v. venusta Trachemys venusta panamensis torals, abdominals, and femorals they often contact one or both ssp. nov. are covered with a swirling pattern seams. In T. v. iversoni the prima- Order Testudines Linnaeus, 1758. reaching the lateral extremities of ry vertebral scute markings are Suborder Cryptodira Cope, 1868. the humeral/pectoral (sometimes often aligned in a parallel fashion Family Emydidae Rafinesque, 1815. pectoral/abdominal), abdominal/ with the median dorsal stripe; not Holotype (designated herein): femoral, and femoral/anal seams. parallel in T. v. venusta. The first FLMNH 52511, juvenile preserved The lateral pattern on the humer- vertebral scute is longer than wide in alcohol, collected in July 1968 by al/pectoral seams is oriented poste- (sometimes equal) in T. v. iversoni; H. W. Campbell in Panama (Canal riorly and is predominantly on the wider than long in T. v. venusta. Zone) Chiva-Chiva Road, 1 km pectoral scute. The plastral pattern The carapace length of T. v. iversoni from Gaillané Highway.

REPTILIA 46 Trachemys venusta panamensis ssp. nov. holotype: note the vertebral scute patterns. Photos: G. Salmon

Paratype (designated herein): Description going laterally, crossing the para- FLMNH 52512, juvenile preserved In T. v. panamensis the postor- median stripes to join the primary in alcohol, collected in July 1968 by bital stripe contacts the orbit, orbitocervical stripe; the mandibu- H. W. Campbell in Panama (Canal widens as it passes over the tympa- lar tomium is finely serrate; pale Zone) Chiva-Chiva Road, 1 km num, then narrows again going yellow (see LEGLER, 1990) pos- from Gaillané Highway. posteriorly; the primary orbitocer- terocentrally located, usually com- Type locality: Chiva-Chiva vical stripe is the same width as plete primary costal ocelli sur- Road (trail), 1 km from Gaillané most of the postorbital stripe, such round light-orange-centered dark (Gaillard) Highway (Fort Clayton that, with the expanded width over areas, and contact the posterior entrance), north of Miraflores the tympanum, the postorbital scute seams; secondary costal Lake, Pacific-side Panama Canal stripe is dominant; the central sec- markings fill the remaining anteri- Zone, Panamá Province, Panama. ondary orbitocervical stripe gradu- or, lateral, and medial surfaces Distribution: “Pacific side of ally becomes as wide going poste- around the primary costal ocelli; Panama from the Azuero Peninsula riorly as the postorbital and prima- light-orange-centered dark areas at least to the Isthmian region” ry orbitocervical stripes; the cover the intermarginal seams; the (LEGLER, 1990, as the “Central mandibular stripe is isolated; the first and fifth vertebral scutes have American Pacific” population). symphyseal stripe does not con- primary markings parallel to the Etymology: As the only known nect with the variable paramedian median line; the second and third endemic form of Trachemys, neck stripes, so no Y is present on vertebral scutes have a central named for the country, Panama, the chin; the median neck stripe on posteriorly open horseshoe design where the turtle naturally occurs. the posterior ventral neck splits (often faded); the second, third,

Left: Distributions of Trachemys taxa as recognized before this paper: Trachemys ornata (purple); Trachemys emolli (red); Trachemys venusta venusta (forest green); Trachemys venusta cataspila (brown); Trachemys venusta grayi (light blue). Right: Distributions of the herein revised Trachemys taxa: Trachemys venusta venusta (orange); Trachemys venusta iversoni ssp. nov. (yellow, with upper red dot indicating Buctzotz and lower red dot Izamal); Trachemys venusta uhrigi ssp. nov. (bright green: confirmed range in Honduras; forest green: probable range; dark green: suspected range; red dot: San Pedro Sula); Trachemys venusta panamensis ssp. nov. (blue, with red dot indicating Chiva-Chiva Rd).

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and especially the fourth vertebral as it goes over the tympanum mak- being dominant. The postorbital scutes have a relatively intricate ing it the dominant lateral head (excluding over the tympanum), pattern of lateral markings; the stripe; in T. v. venusta it is consis- primary orbitocervical, and posteri- carapace of juveniles is widest at tantly thin, with the primary orbito- or central secondary orbitocervical the seam between the 7th and 8th cervical stripe being dominant. stripes are all the same moderate marginals; first vertebral scute is The central secondary orbitocervi- width in T. v. panamensis; in T. v. wider than long; the bridges have cal stripe in T. v. panamensis uhrigi the postorbital and central two light lines each; the plastral widens to match that of the postor- secondary orbitocervical stripes are pattern of hatchlings is reduced bital and primary orbitocervical thin and the primary orbitocervical with bold dark lines longitudinally going posteriorly; thin in T. v. stripe is moderate in width. On the following both sides of the median venusta. On the lower ventral neck lower ventral neck of T. v. pana- scute seams as far as the posterior of T. v. panamensis the median mensis the median stripe splits lat- half of the intergular seam and the stripe splits laterally and joins the erally and joins the primary orbito- anterior half of the interanal seam. primary orbitocervical stripe; not cervical stripes; instead, in T. v. Laterally the bold lines expand to so in T. v. venusta. uhrigi there is often a transverse involve 90% of the gular/humeral Carapace. The primary carapa- yellow stripe only on the upper seams, 20–30% of the humeral/ cial scute markings are pale yellow- ventral neck joining the paramedi- pectoral seams, 80–90% of the pec- orange in T. v. panamensis; bold an stripes at or near the anterior toral/abdominal seams, 95–100% orange in T. v. venusta. The bolder median stripe, creating an H-like of the abdominal/femoral seams, vertebral scute markings of T. v. figure. and 100% of the femoral/anal panamensis vary from scute to Carapace. The primary costal seams. In total 30–40% of the scute; consistently parallel to the ocelli in T. v. panamensis juveniles plastron is covered by the pattern dorsal midline in T. v. venusta. The are complete as they meet the pos- in hatchlings. The plastral seam carapace of T. v. panamensis is terior scute seam and have formula in hatchlings likely starts widest at the seam between the 7th orange-centered central dark with abd > an > pect. The central and 8th marginals; in T. v. venusta areas; in T. v. uhrigi juveniles stripe on the rump appears to be at the 7th marginal. these ocelli are incomplete poste- solid. Plastron. The plastral pattern of riorly, and have green-centered T. v. panamensis juveniles is central dark areas. The bolder Diagnosis reduced, occupying 30–40% of the vertebral scute markings of T. v. We herein differentiate T. v. plastron; in T. v. venusta it is panamensis are more variable than panamensis ssp. nov. from the greatly reduced and occupies 20% in T. v. uhrigi. nominotypical (conspecific) form of the plastron. In juvenile T. v. Plastron. In T. v. panamensis T. v. venusta sensu stricto (as de- panamensis the interpectoral seam each bridge has two light lines; scribed with the range given above) is the third largest seam; in T. v. three or four in T. v. uhrigi. The and its geographically closest conspe- venusta the third largest is the plastral pattern of juvenile T. v. cific form T. v. uhrigi. Of the charac- interfemoral or intergular. panamensis is reduced, occupying ters given in the foregoing descrip- Trachemys v. panamensis can be 30–40% of the plastron; in T. v. tions, only those differentiating T. v. differentiated from T. v. uhrigi as uhrigi it is greatly expanded, occu- panamensis from T. v. venusta and follows: pying 90–95% of the plastron. The T. v. uhrigi are given here. Head and neck. In T. v. pana- plastral pattern of juvenile T. v. Trachemys v. panamensis can be mensis the postorbital stripe panamensis never reaches the pos- differentiated from T. v. venusta as widens over the tympanum making terior end of the interanal seam; it follows: it the dominant head stripe; in T. v. does in T. v. uhrigi juveniles. In T. Head and neck. In T. v. pana- uhrigi it is consistently thin, with v. panamensis juveniles the inter- mensis the postorbital stripe widens the primary orbitocervical stripe pectoral seam is usually the third

CL CW6 CW7 CW8 CD V1L V1W PL BL IG IH IP IAb IF IAn HL HW HD Trachemys venusta uhrigi (FLNHM) #157800 holotype 261.91 184.70 188.54 186.99 92.95 43.64 40.81 215.92 89.31 30.39 22.94 28.08 63.76 31.22 37.41 58.76 39.19 31.53 (FLNHM) #105425 paratype 135.39 101.54 107.92 105.58 49.41 23.91 29.02 123.48 48.38 20.03 9.89 21.03 28.45 17.35 24.75 31.09 21.86 17.22 Trachemys venusta iversoni (FLNHM) #50478 holotype 187.07 139.55 139.28 135.80 75.31 33.57 26.92 165.96 66.27 24.73 12.85 24.60 43.62 27.56 31.32 44.47 31.46 23.53 (FLNHM) #50476 paratype 126.26 92.98 96.70 94.25 54.36 23.92 23.08 111.64 47.54 17.32 8.43 20.78 29.41 13.95 21.34 31.66 23.88 16.92 Trachemys venusta panamensis (FLNHM) #52511 holotype 40.09 37.36 37.67 36.31 15.52 9.52 12.07 35.34 11.91 7.53 2.41 6.21 5.67 5.75 7.75 13.74 10.18 6.16 (FLNHM) #52512 paratype 38.42 34.00 34.11 34.19 15.67 8.96 10.84 34.51 10.93 6.50 2.97 5.53 7.00 4.51 7.88 12.23 9.75 6.66

Table 1. Measurement data (in mm) for three new subspecies of Trachemys venusta. Morphometric key: CL = straight midline carapace length; CW 6, 7, 8 = straight carapace width at 6th, 7th and 8th marginals; CD = maximum carapace depth; V1L = first vertebral length; V1W = first vertebral width; PL = midline plastral length; BL = bridge length; IG, IH, IP, IAb, IF, IAn = midline plastral inter-scute seam lengths; HL = maximum head length; HW = head width at tympana; HD = maximum head depth.

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longest plastral seam; in T. v. uhrigi the third longest is the femoral. The central stripe on the rump of T. v. panamensis is usually solid; bro- ken in T. v. uhrigi. ■

Acknowledgments We thank Dennis Uhrig for access to his vast collection of Trachemys Trachemys emolli Trachemys ornata and his many helpful comments; Roger Bour, John Iverson, Gerard Salmon, and Steven Winchell for constructive criticism; Kraig Adler, John Iverson, and Ned Gilmore for rare literature; David Kizirian and Rob Pascocello (AMNH), Ned Gilmore (ANSP), Colin McCarthy (BMNH), Kenneth L. Krysko and Mel Gramke (FLMNH), and Trachemys venusta venusta sensu stricto Trachemys venusta uhrigi ssp.nov. Stephan Böhm for facilitating access to specimens; and Kenneth Krysko and Mel Gramke for speci- men accession at the Florida Museum of Natural History. Manuscript preparation was partial- ly supported by the Department of Energy under Award Number DE- FC-09-075R22506 to the University Trachemys venusta grayi Trachemys venusta iversoni ssp.nov. of Georgia Research Foundation.

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