TU' ^^^(^,c^ HARVARD UNIVERSITY Library of the Museum of Comparative Zoology zz^ VOLUME I 1980-1981 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY, a publication of the Biology Department of Tulane University, is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on areas outside this geographic area will be considered. Each number contains an indivi- dual monographic study or several minor studies. Normally two numbers plus an index and a table of contents are issued annually. Preferred citation of the journal is Tulane Stud. Zool. and Bot. INFORMATION FOR AUTHORS: Manuscripts submitted for publications are eval- uated by the editors and by an editorial committee selected for each paper. Contrib- utors need not be members of the Tulane faculty. Manuscripts of 20 or more pages, double-spaced, are preferred. We recommend conformance with the principles stated in CBE Style Manual, 4th ed., 1978. Manuscripts should be typewritten and double spaced. Two additional copies should accompany the original to expedite editing and publication. Legends for figures should appear on a separate page and in sequence. Illustrations should be proportioned for one or two column width corresponding to our printed page size, and should allow for insertion of the legend if occupying a whole page. Guidelines for letter and other extraneous markings should be done with a non-photo blue pencil such as Eagle Prismacolor. Photographs should be on glossy paper. Many tables, if carefully prepared with a carbon ribbon and electric typewriter, can be photographically reproduced, thus helping to reduce publication costs. Lettering in any illustrative or tabular material should be of such a size that no letter will be less than 1 Vi mm high when reduced for publication. An abstract not exceeding three percent of the length of the article must accompany the manuscript. Separates of published articles are available to authors at a nominal cost. Page charges, calculated at $45/page, are solicited from authors who have funds for this purpose through their institutions or grants. Acceptance of papers is not dependent on ability to underwrite costs but excessive illustrations and tabular matter may be charged to the author. EXCHANGES, SUBSCRIPTIONS, ORDERS FOR INDIVIDUAL COPIES: Ex- changes are invited from institutions publishing comparable series. Subscriptions are billed in advance. A price list of back issues is available on request. Individuals should send their remittance, preferably money order, along with their orders. Remittances should be made payable to "Tulane University." Subscription rates: Volume 21, 22 $7.50 ea., $8.50 foreign. Volume 23. $8.50 domestic, $9.50 foreign. Copies of Tulane Studies in Zoology and Botany sent to regular recipients, if lost in the mails, will be replaced if the editorial offices are notified before the second subsequent issue is released. COMMUNICATIONS: Address all queries and orders to: Editor, TSZ&B, Depart- ment of Biology, Tulane University, New Orleans, Louisiana 701 18, U.S.A. Harold A. Dundee, Editor Arthur L. Welden, Associate Editor Samuel Clifford, Assistant to the Editors CONTENTS OF VOLUME 22 Number Page 1. THE SHREVEI GROUP OE HISPANIOLAN SPHAERODACTY- LUS (REPTILIA, GEKKONIDAE) Albert Schwartz and Eugene D. Graham, Jr. 1 NATURAL HISTORY OF THE MAP TURTLES GRAPTEMYS PSEUDOGEOGRAPHICA and G. OUACHITENSIS IN WISCON- SIN Richard C.Vogt 17 A TAXONOMIC STUDY OF THE COOTER TURTLES PSEU- DEMYS FLORIDANA (LECONTE) AND PSEUDEMYS CON- CINNA (LECONTE) IN THE LOWER RED RIVER, ATCHAFA- LAYA RIVER, AND MISSISSIPPI RIVER BASINS Kenneth Mark Fahey 49 2. LIFE HISTORY PATTERNS OF NOTROPIS SABINAE (PISCES: CYPRINIDAE) IN THE LOWER SABINE RIVER DRAINAGE OF LOUISIANA AND TEXAS David C. Heins 67 THE SYSTEMATICS AND DISTRIBUTION OF THE HOGNOSE VIPER BOTH RO PS NASUTA BOCOURT (SERPENTES: VIPERI- DAE) Louis Porras, James L. McCranie, and Larry David Wilson 85 SOME TREMATODES OF MAMMALS IN LOUISIANA Wesley L. Shoop and Kenneth C. Corkum 109 ISSN 0082-6782 JUN4 1980 f- •; .'N P^ \/ /A , T O Volume 22, Number 1 $4.00 'May 26, TY THE .SAy/?EV^E/ GROUP OF HISPANIOLAN 5P///^E/?OD/4CryLty5 (REPTILIA, GEKKONIDAE) ALBERT SCHWARTZ and EUGENE D. GRAHAM. JR. p. 1 NATURAL HISTORY OF THE MAP TURTLES GRAPTEMYS PSEUDOGEOGRAPHICA AND G. OUACHITENSIS IN WISCONSIN RICHARD C. VOGT p. 17 A TAXONOMIC STUDY OF THE COOTER TURTLES, PSEUDEMYS ELORIDANA (LECONTE) AND PSEUDEMYS CONCINNA (LECONTE), IN THE LOWER RED RIVER. ATCHAFALAYA RIVER. AND MISSISSIPPI RIVER BASINS KENNETH MARK FAHEY P- 49 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY, a publication of the Biology Department of Tulane University, is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on areas outside this geographic area will be considered. Each number contains an indivi- dual monographic study or several minor studies. Normally two numbers plus an index and a table of contents are issued annually. Preferred citation of the journal is Tulane Stud. Zool. and Bot. INFORMATION FOR AUTHORS: Manuscripts submitted for publications are eval- uated by the editors and by an editorial committee selected for each paper. Contrib- utors need not be members of the Tulane faculty. Manuscripts of 20 or more pages, double-spaced, are preferred. We recommend conformance with the principles stated in CBE Style Manual, 4th ed., 1978. Manuscripts should be typewritten and double spaced. Two additional copies should accompany the original to expedite editing and publication. Legends for figures should appear on a separate page and in sequence. Illustrations should be proportioned for one or two column width corresponding to our printed page size, and should allow for insertion of the legend if occupying a whole page. Guidelines for letter and other extraneous markings should be done with a non-photo blue pencil such as Eagle Prismacolor. Photographs should be on glossy paper. Many tables, if carefully prepared with a carbon ribbon and electric typewriter, can be photographically reproduced, thus helping to reduce publication costs. Lettering in any illustrative or tabular material should be of such a size that no letter will be less than 1 V2 mm high when reduced for publication. An abstract not exceeding three percent of the length of the article must accompany the manuscript. Separates of published articles are available to authors at a nominal cost. Page charges, calculated at $45/page, are solicited from authors who have funds for this purpose through their institutions or grants. Acceptance of papers is not dependent on ability to underwrite costs but excessive illustrations and tabular matter may be charged to the author. EXCHANGES, SUBSCRIPTIONS, ORDERS FOR INDIVIDUAL COPIES: Ex- changes are invited from institutions publishing comparable series. Subscriptions are billed in advance. A price list of back issues is available on request. Individuals should send their remittance, preferably money order, along with their orders. Remittances should be made payable to "Tulane University.'' Subscription rates: Volume 22 $7.50, $8.50 foreign. Copies of Tulane Studies in Zoology and Botany sent to regular recipients, if lost in the mails, will be replaced if the editorial offices are notified before the second subsequent issue is released. COMMUNICATIONS: Address all queries and orders to: Editor, TSZ&B, Depart- ment of Biology, Tulane University, New Orleans, Louisiana 701 18, U.S.A. Harold A. Dundee, Editor Arthur L. Welden, Associate Editor Samuel Clifford, Assistant to the Editors TULANE STUDIES IN ZOOLOGY AND BOTANY Volume 22, Number 1 May 26, 1980 THE SHREVEI GROUP OF HISPANIOLAN SPHAERODACTYLUS (REPTILIA, GEKKONIDAE) ALBERT SCHWARTZ and EUGENE D. GRAHAM, JR. Miami-Dade Community College North, Miami, Florida 33167 and 717 S.W. 7th St., Hallandale, Florida 33009 ABSTRACT The Hispaniolan shrevei group of Sphaerodac- tyius is composed of four species, one of which (S. asterulus) is described in the present paper. Detailed descriptions of the four included species are presented, as well as a hypothetical history of the group. S. shrevei stands alone, whereas the other species show an interrelationship in pattern. The group is considered to be not closely related to other Hispaniolan or Antillean Sphaerodactylus and to be north island (sensu Wilhams) in affinities. On the Antillean island of Hispaniola occurs a group of species of Sphaerodac- tylus without obvious close relatives there or elsewhere in the West Indies. The first- named of this group of lizards is S. shrevei Lazell, and the group-name is derived from this species. Two other species have been described: S. rhabdotus Schwartz and 5. leucaster Schwartz. The latter two species were named on limited material; there was only the single female holotype of S. shrevei. Sphaerodactylus rhabdotus and 5. leucaster have complementary dis- tributions in the Dominican Valle de Neiba and the contiguous Llanos de Azua to the east, and 5. leucaster extends northward toward the Valle de San Juan. Sphaerodactylus shrevei, on the other hand, was described from the Haitian Presqu'ile du Nord Ouest, at Mole St. Nicholas, removed some 250 km airline from the nearest records for S. rhabdo- tus. We have re-examined the holotype (formerly lost but now found) in light of six new specimens. Small primary series of S. leucaster and S. rhabdotus have gradually been amplified with additional specimens (rarely series), so that we now know more about the variation in these two species than we did previously. One problem in obtaining examples of this group of sphaerodactyls is that they are xerophiles, inhabiting rigorous and often hazardous terrain - cactus and Acacia forest - where collecting is dif- ficult. Other than the very large series of the new species described herein, most of which was native-collected, the senior au- thor secured only one specimen from na- tive collectors. The edificarian situations that allowed the collection of the original lots of 5. leucaster and S. rhabdotus no longer exist. Search for these two species of geckos in non-edificarian situations under fallen and decayed arborescent cactus trunks, in decayed Acacia trunks, both prostrate and standing, in palm lit- ter, and under rocks, as well as under and in the dead basal rosettes oi Agave leaves has yielded occasional specimens. This labor has resulted in the accumulation of EDITORIAL COMMITTEE FOR THIS PAPER: DR. ERNEST E. WILLIAMS, Curator of Reptiles and Amphibians, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138 DR. RICHARD THOMAS, Assistant Professor of Biology, University of Puerto Rico, Rio Piedras, Puerto Rico 00931 1 Tulane Studies in Zoology and Botany Vol. 22 i 19 specimens of S. leucaster and 27 of S. rhabdotiis. The largest single series of ei- ther species is 15 specimens oiS. rhabdo- tiis from the type-locality, with nine specimens of the same species collected by James R. Dixon near La Descubierta. The series of eight specimens of S. rhab- dotiis from the type-locality is the largest series of that species; all other localities are represented by one or two specimens. We emphasize the above for a particu- lar reason. In August 1977, we had brought to us, with many specimens of S. cinereiis Wagler, at Terre Sonnain, north of Les Poteaux, a single specimen of a gecko that obviously belonged to the shrevei group. The lizard was in poor con- dition. It seemed likely that we were at the periphery of its range, since among many lizards, this one specimen was brought to us. Another possibility was that the native collectors there were col- lecting in a fashion that did not encourage the taking of this type of gecko. Since S. cinereus is primarily edificarian or arbor- eal (under the loose bark of trees), it is conceivable that the local Haitians were concentrating their efforts in such situa- tions and were not looking under ground- cover as a general collecting procedure. On 20 June 1978, we traveled the coastal road between Gonaives and Coridon for a few kilometers west of ^a Soleil. This trip, west of the Terre Sonnain locality, was an effort to locate more individuals of the same species. We stopped at an uninhabited locality 9.3 km W (;a Soleil (Fig. 3). The road at this point passes through a series of lime- stone hills with cliffs, many loose rocks, boulders, and other debris; the most con- spicuous plant is Agave, of which there were not only large living plants but many decaying basal rosettes of leaves of plants long dead. In addition,' there were small piles of rocks scattered over a relatively level area between two hills. Everything seemed to be appropriate for finding some sort of xerophilic gecko. We had success almost immediately and secured a series of 42 lizards in about 45 minutes by searching in the rock piles and in the Agave rosettes; five unhatched eggs (and many empty shells) were also found. The series of lizards agreed with our single specimen from Terre Sonnain from the previous summer. Most amazingly, this species appeared to be locally extremely abundant in contrast to all other mem- bers of the group. As we traveled beyond this locality, we encountered a few roadside huts as well as a small village whose name, after that of this entire region, is Lapierre. Inhabit- ants of this village were avid collectors and obtained an immense series for us. We later proceeded across the limestone hills to a sea level oasis, where this gecko was also abundant (although it occurred there with two other species, one similar to S. altavelensis Noble and Hassler and the other undescribed). We have only one record from farther along (15.7 km W fa Soleil) the road to Coridon, but we feel certain that its distribution continues still farther along the very xeric and very sparsely inhabited coast. At the town of Coridon, we were unsuccessful in secur- ing specimens but feel that this was due to the abundance there of the much more easily secured Ameiva chrysolaema Cope - the only lizard brought to us by the local inhabitants. Considering the proximity of the Lapi- erre area to the type-locality of 5. shrevei (80 km airline) and its distance from the nearest record of S. rhabdotiis (130 km airline), we were struck by the similarity in coloration and pattern of these new lizards to S. rhabdotiis and S. leucaster, and not to S. shrevei. Basically, S. shrevei is blotched, whereas S. leucaster and S. rhabdotiis are longitudinally striped or lined. The new geckos agree with the lat- No. 1 Hispaniolan Sphaerodactylus ter pair, most especially with 5. leucaster in that the longitudinal lines are inter- rupted by tiny white dots. These dots are much smaller than are the similar white blotches or "stars" that interrupt the lon- gitudinal lines in S. leucaster. We accu- mulated a series of 448 specimens of the new species from the Lapierre region. With this huge lot of specimens, as well as the more abundant material of both S. leucaster and S. rhabdotus, this seems an appropriate time to review this group of xerophilic geckos. The shrevei group may be character- ized by the following combination of characters: 1) size moderate; maximum size 31 mm snout-vent length in males, 32 mm in females; 2) dorsal scales large, keeled, subimbricate to not imbricate, and with or without a few scattered mid- dorsal smaller scales forming an incom- plete middorsal zone; 3) dorsal scales with dorsal smaller scales forming an incomp- lete middorsal zone; 3) dorsal scales with hair-bearing scale organs only; 4) throat, chest, and ventral scales smooth; 5) in- ternasal scales usually 2; 6) supralabial scales to mid-eye usually 4; 7) escutcheon extremely variable within species, from 1 to 9 scales in length and from 6 to 25 scales in width, maximally extending to behind the knees laterally; 8) sexual di- chromatism weak to absent in basic pat- tern, but coloration of head different in males and females; 9) dorsal pattern ei- ther lineate or blotched, head in males yellow to orange, no dark head pattern present; 10) no scapular patch and in- cluded ocelli; 11) ontogenetic variation in dorsal pattern apparently absent; 12) head short and blunt, with a convex snout; 13) pupil of eye vertically elliptical rather than circular. TAXONOMIC ACCOUNTS Sphaerodactylus shrevei Lazell Sphaerodactylus shrevei Lazell, 1961, Breviora(139):l. Type-locality: Mole St. Nicholas, Departement du Nord Quest, Haiti. Holotype: MCZ 62548. Definition. An Hispaniolan species of Sphaerodactylus characterized by the combination of: 1) dorsal scales subim- bricate or not imbricate, with an ill de- fined middorsal zone of smaller keeled scales not forming a continuous dorsal row; 2) high number of scales (38-46) around midbody; 3) modally 1 intemasal scale; 4) dorsal pattern basically blotched and not lineate, with about 5, more or less symmetrical, pairs of blotches on each side of the dorsal midhne (Fig. 2); 5) an irregular ventrolateral line of much smal- ler blotch-like figures. Description. There are now seven specimens of 5. shrevei: the holotypic fe- male, three adult females, one subadult male, and 2 juveniles (snout-vent lengths 14 and 17 mm). Scale counts and mea- surements for the series are: largest male (ASFS V49555) with a snout- vent length of 24 mm, largest females (MCZ 62548, ASFS V49769, ASFS V49831) 30 mm; dorsal scales between axilla and groin 22- 27 (x = 23.8); ventral scales between ax- illa and groin 31-33 (32.2); midbody scales 38-46 (42.0); supralabials to mid- eye modally 4/4 (five individuals) but two individuals have counts of 5/5; intemasal modally 1 (5 individuals), but two speci- mens have 2 internasals; fourth toe lamel- lae 10-11 (10.3; Mq = 10 - 75%); escutch- eon in single male 6 X 25, consisting of a central group of thickened scales with a single row of scales extending to behind the knee. Lazell's description of the holo- type agrees well with the condition in the fresh material. Sphaerodactylus shrevei is a blotched (rather than lineate) lizard. There is a broad dark gray cephalic U enclosing a darkly stippled snout, fol- lowed by a supraaxillary pair and a post- axillary pair of more or less rectangular Tulane Studies in Zoqlogy and Botany Vol. 21} dark blotches; these are in turn followed by about five irregular crossbands that are often broken middorsally to give a series of lateral dark blotches. The ground color is gray dorsally and white ventrally. The two juveniles do not differ markedly from the adults as far as colora- tion or pattern. Lazell described the female holotype in life as follows: "a dull-colored animal with a pattern composed of three basic hues - each tending to be unique on an Remarks. S. shrevei is so very distinc- tive in having a blotched pattern that we are not really sure that it is at all closely related to the balance of the species in this group. Although, as will be shown later, some S. leucaster approach blotch- ing to some extent, they do not have such a markedly blotched pattern as does S. shrevei. Comparisons with other taxa will be made in their appropriate places in the present paper. Lazell (1961) stated that the holotype individual scale. There are very irregular was taken from a large circular rock pile. dark gray-brown blotches across the dor- sum; beginning at the back of the head there are three such markings to the shoulders. There are three more crudely 'Y' shaped markings on the body, the most anterior of which bifurcates to the right, the remaining ones bifurcating to the left. There are two small blotches on the right side of the rump and one on the left. The second transverse blotch, on the nape, is broken by a light middorsal line that continues down through the fourth V49831). marking and then fades out. The ground color of the dorsum is ash gray. There are scattered over the dorsal surface short transverse series of white or partly white scales - from two to four in a row - that appear to have no correlation whatever with the rest of the animal's pattern. The top of the head is ash gray except for a very irregular, dark, gray-brown blotch on the parietal area. Coming back from the eye are two stripes, one of which runs downward across the cheek; the other nearly connects with the first transverse marking on the back of the head . . . Along each side of the animal is a line of partially connected, small, dark, gray- brown blotches; just ventral to this row is another composed of widely spaced, single dark, gray-brown scales." the rocks varying in size from golf ball to nearly football size. The pile was of some age since several Acacia were growing from within it. The area at M3le St. Nich- olas is arid. The new material was native- collected and we have no data on the habitat. Specimen exumined. Haiti, Depi. du Nord Quest, Mole St. Nicholas (MCZ 62548 - holotype; ASFS V49555-56, ASFS V49586-87, ASFS V49769, ASFS Distribution: Known only from the type-locality (Fig. 1). Sphaerodactylus rhabdotus Schwartz Sphaerodactylus rhabdotus Schwartz, 1970, J. Herpetology, 4(l-2):64. Type-locality: 5 km SE La Florida, 500 feet (153 m), Independencia Province, Republica Dominicana. Holotype: USNM 166960. Definition. An Hispaniolan species of Sphaerodactylus characterized by the combination of: 1) dorsal scales keeled and only slightly imbricate, a few small keeled scales in the shoulder region but no middorsal row of granules or small scales; 2) moderate number (36-44) of scales at midbody; 3) modally 2 internasal scales; 4) dorsal pattern of two dark brown lines on a pale ground, these lines not interrupted by white dots or spots; a brownish occipital Y usually present (Fig. 2). No. 1 His pan iolan Sphaerodactylus Distribution. Republica Dominicana, Independencia Province, from near Las Lajas on the Dominico-Haitian border, east to the vicinity of Angostura, and oc- curring on both the north and south sides of Lago Enriquillo (Fig. 1). Variation. There are 27 specimens of S. rhahdotus; the largest male (USNM 166960 - holotype) has a snout-vent length of 31 mm, the largest female (ASFS V20486) 32 mm. Scale counts on the series are: dorsal scales between ax- illa and groin 20-28 (x = 23.8); ventral scales between axilla and groin 24-31 (28.0); midbody scales 36-44 (40.4); su- pralabials to mid-eye modally 4/4 (23 in- dividuals) but other counts of 4/5 (2), 4/6 ( 1 ), and 5/5 (1); internasals modally 2 (17 individuals), but other counts of 1 (2) or 3 (8); fourth toe lamellae 9-12 (10. 1; Mq = 9 - 36%); escutcheon 5-9 X 7-25. In life, the dorsal ground color varies from pink (almost lavender) to very pale gray; the dorsolateral lines are brown to black. Fig. 1. Hispaniola, eastern Haiti and western Re- publica Dominicana, showing the known distribu- tions of Sphaerodactylus of the shrevei group: hexa- gon, 5. shrevei; squares, S. asterulus; circles, S. rhahdotus; triangles, S. leucaster. Some localities for S. asterulus have been omitted for clarity. these lines usually entire and without in- terruption (but see beyond). An occipital Y is present but variable in expression (i.e. , from clear and distinct to somewhat blurred); the stem of the Y may lead to a median dorsal brown line so that the dor- sum appears trilineate. Males have the head and snout yellow in life, and adults of both sexes have the tail yellow; this latter feature is more pronounced in males than in females. The iris has been recorded as brown with a golden pupil- lary ring, or gray. There may be a few irregularly placed middorsal small, non- granular scales, especially in the shoulder region at the zone of transition from the granular nuchal scales and the large dor- sals. Comparisons. As far as size and scutel- lation are concerned, the differences bet- ween S. shrevei and S. rhahdotus are slight. Females reach about the same snout-vent lengths (30 mm in S. shrevei, 32 mm in S. rhahdotus). Means of dorsals in axilla-groin distance are the same (23.8). However, 5. shrevei averages higher in number of ventrals (32.2 versus 28.0) and midbody scales (42.0 versus 40.4). In the latter count, the scale counts of the two species overlap broadly, whereas in the former the counts overlap but slightly (31-33 in S. shrevei, 24-31 in S. rhahdotus). The single internasal in S. shrevei differs from the 2 internasals in S. rhahdotus; the modes in each case are strong. But the two species differ amply in dorsal pattern and coloration; S. shre- vei is a blotched lizard and lacks an occipi- tal Y, whereas S. rhahdotus is lineate and has an occipital Y. The hues involved in both cases are distinctive. S. rhahdotus is a pallid lizard, dorsally striped with brown on a pale pink to lavender ground, in contrast to the gray dorsal color and blotched pattern of S. shrevei. The dorsal pattern of S. rhahdotus is quite constant. But there is one specimen Tulane Studies in Zoology and Botany Vol. 22 \ (TCWC) that requires comment. This in- dividual (male, snout-venth length 26 mm) has the dorsolateral brown lines so broken as to cause them to form a series of dots. Although not all specimens of S. rhabdotus have the lines absolutely com- plete, this individual is at the extreme condition. Faded specimens of 5. leucas- ter, which has the dorsolateral lines inter- rupted by (in life) white, starlike blotch- es, assume this same pattern (i.e., the white pigmentation fades leaving only broken dark lines). This specimen (TCWC 50162) thus superficially resem- bles preserved specimens of S. leucaster; whether in life it had white spots in the breaks in the dorsolateral stripes is un- known. If so. it would certainly re-enforce the relationship of S. rhabdotus and geo- graphically close S. leucaster and would suggest that these two species are in actu- ality subspecies. At present their ranges are known to be complementary, but they differ in dorsal pattern. Perhaps these differences are at the subspecific level, but we are unwilling as yet to take that taxonomic step. The type series was collected in the fallen thatch of a construction-crew shack at the northern base of the Sierra de Ba- oruco. The lizards were relatively abun- dant in the thatch, especially at the in- terface between the thatch and the lime- stone base upon which it was resting, and within the dry, folded, accordion-like bases of the palm fronds. The specimen from near Angostura was taken from be- neath a piece of dry palm siding at an old and abandoned shack near a gypsum mine. The Duverge' lizard was found in an old pile of palm trunk cuttings in xeric scrub. The senior author has never col- lected S. rhabdotus in a "natural" situa- tion, despite search for it in the xeric Valle de Neiba. Specimens examined. Republica Do- minicana, Independencia Province, 1 km ESE Las Lajas, 75 m (UF/FSM 42920); 5 km SE La Florida, 153 m (USNM 166960 - holotype, USNM 166961-62, CM 45890-92, MCZ 92096- 98, ASFS V17180, ASFS V20485-87, ASFS V20797-98); 6 km NW Duverge (RT 4421); 2.0 km E La Descubierta, 15 m (TCWC 50161-69); 11 km SE Ango- stura (ASFS V4 1298). Sphaerodactyhis leucaster Schwartz Sphaerodactylus leucaster Schwartz. 1973, Proc. Biol. Soc. Washington, 86 (4):36. ry/7e-/oca//rv.- EI Iguito, 1.6 mi. (2.6 km) NE Fondo Negro, Barahona Province, Republica Dominicana. Ho- lotype: USNM 189234. Definition. An Hispaniolan species of Sphaerodactylus characterized by the combination of: 1) dorsal scales keeled and slightly imbricate, with a few smaller scaled middorsally; 2) moderate number (35-46) of scales at midbody ; 3) modally 2 internasal scales; 4) dorsal pattern of 2 dark brown dorsolateral and 2 broken ventrolateral stripes on a lavender ground, the dorsolateral stripes with from 4 to 7, relatively large, discrete white spots per stripe; 5) males with a median dark brown nuchal line, females with a dark brown cephalic-nuchal Y (Fig. 2); heads and throats bright orange and tails yellow in males. Distribution. Republica Dominicana, the extreme eastern portion of the Valle de Neiba, extending thence eastward as far as the vicinity of Bani, Peravia Pro- vince, in the Llanos de Azua (Fig. 1). Variation. There are 19 specimesn of S. leucaster; the largest male (ASFS V40797) has a snout-vent length of 27 mm, the largest female (ASFS V35639) 31 mm. Scale counts on the series are: dorsals between axilla and groin 19-29 (15 = 23.5); ventral scales between axilla and groin 20-32 (27.4); midbody scales 35-46 (39.8); supralabials to mid-eye 4/4 (15 » No. 1 Hispatiiolan Sphacrodactylus individuals), 3/3 (2), and 5/5 (2); inter- nasals modally 2 (13 individuals) or 1 (6); fourth toe lamellae 7-11 (9.5; M^ = 10 - 44 ^r); escutcheon 5-8 X 8-24. The dorsal color in life is lavender, with, in males. pecies. The two closest localities for the species are 11 km SE Angostura, Inde- pendencia Province, and 2 km NE Can- oa. Barahona Province - a distance of 21 km. The presence of a single specimen the head bright orange; there is a pair of of .V. rhahdotus from near La Descubierta dorsolateral dark brown lines that begin having a pattern similar to that of 5. leu- behind the eyes and extend onto the base caster suggests that there is genetic con- of the tail. Ventral to these, there is a pair tinuity between the two taxa. But this is of lateral dark brown lines. The dorsolat- eral lines in life are broken by a series of distinct white blotches; in preserved specimens these white blotches disappear (due in part at least to the general pale dorsal color) with the result that the dor- solateral lines appear merely broken. The venter is flesh-colored, and the tail yellow in males. Some specimens have a vague median nuchal line which may lead to a pair of arms extending onto the head, to form an occipital Y. Females are pat- terned like males except that the head is not orange, and there may be some dark brown stippling on the throat. There are a few scattered smaller scales middorsally. Comparisons. S. leucaster most closely resembles S. rhabdotus in both coloration and Uneate pattern. The major pattern difference between these two species is the presence of the white spots in the dorsolateral lines of S. leucaster to give a broken effect. From S. shrevei, S. leucas- ter differs in being lineate and not blotched (but see below). Scale counts between S. leucaster and S. rhabdotus are comparable. S. shrevei lies at the upper extreme of S. leucaster in midbody scale counts, and the mode in S. shrevei is one internasal (71%) a condition observed in about one-third of the S. leucaster exam- ined. The iris in 5. leucaster is greenish gold, and brown to gray in S. rhabdotus; there are no data on the iris color in S. shrevei. Remarks. We have already comment- ed on the possibility that S. rhabdotus and S. leucaster should be regarded as subs- slim evidence at best, and neither species is represented from many localities or in series. Perhaps the most interesting phenome- non in S. leucaster occurs in three speci- mens from the eastern and northern ex- tremes of the range. Four specimens re- quire comment. ASFS V3579() was brought to the sen- ior author with a long series of S. ocoae Schwartz and Thomas, from north of Cruce de Ocoa. It was not noted as being different in coloration or pattern from that species at the time of its acquisition, and thus no color notes were made on it in Ufe. Only under the microscope was it obvious that this individual was a shrevei group member (and at that time the eas- ternmost specimen). It differs from other S. leucaster in lacking dorsolateral lines and white spots; rather, the dorsum is dotted with dark brown on a brown ground. It is an adult female with a snout- vent length of 29 mm. A second noteworty specimen is UF/ FSM 36964, a female with a snout-vent length of 26 mm. This specimen is from near Corte's in Azua Province, and thus from the northeastern extreme of the range. It is like the specimen described above, although there are distinct rem- nants (much broken) of four dorsal lines with some white flecking still distinct on the head (but not the body). UF/FSM 42926 is a female with a snout-vent length of 28 mm. This speci- men is from 2 km NE Bani, the extreme eastern limit of S. leucaster. The dorsal ■II 8 Tulane Studies in Zoology and Botany Vol. 22 pattern is without any obvious lineate configuration but is rather blotched with large dark blotches, very similar to the condition in S. shrevei. Color notes by Fred G. Thompson state that in life the ground color was light grayish blue, head and tail orange, venter more gray than blue, and diffuse black spots on the dor- sum. All three of these specimens men- tioned lack a nuchal Y. A fourth specimen is less peculiar; it is ASFS V35638, a male with a snout-vent length of 26 mm, from the type-locality. In this individual, the dorsolateral lines are very broad and broken into a series of dark blotches. The occipital Y is present. The lizard is distinctive only in the width of the dorsolateral lines and the approach to a blotched condition, although the blotching is much less obvious than in UF/FSM 42926. The significance of these peculiar var- iants is prolematical. In each case except the last, there is only one specimen from a locality. Whether there are other species (or subspecies of S. leucaster) in the east- ern portion of the area we attribute to S. leucaster remains undetermined. Yet from, or between, some of these locali- ties, there are specimens that are typical S. leucaster. None of the prolematical specimens is maximally sized. The senior author has tried repeatedly to secure ad- ditional specimens at the locality for ASFS V3579() (3 km N Cruce de Ocoa, Peravia Province), without success. In fact, the area is oasis-like in an otherwise very xeric region, not the typical situation for a member of the shrevei group. More likely, the local S. leucaster population occurs in the adjacent desert areas, where it may be difficult to secure additional specimens. Aside from the highly specialized situa- tion at the type-locality, S. leucaster has been taken in an abandoned termitarium and under the bark of a large, standing. old Acacia near the base, both in xeric coastal woods, and under limestone boulders on a xeric hillside. Fred G. Thompson secured specimens in the dead basal rosettes oi Agave. Specimens examined. Republica Do- minicana, Barahona Province, 2 km NE Canoa, 50 m (UF/FSM 42924); 3 km NE Canoa, 110 m (UF/FSM 42921); 13 km ESE Canoa (UF/FSM 42922-23); 15 km ESE Canoa (ASFS V40797-98); El Igui- to, 2.6 km NE Fondo Negro (USNM 189234 - holotype, ASFS V30524-26, ASFS V35637-40); Azua Province, 2 km NW Cortes, 240 m (UF/FSM 36964); 3.5 km SSE Azua, 30 m (UF/FSM 42925); 5.3 km ESE Azua (USNM 197338); Per- avia Province, 3 km N Cruce de Ocoa, 153 m (ASFS V35790); 2 km NE BanC 100 m (UF/FSM 42926). Sphaerodactylus asterulus, new species Holotype. CM 60521, an adult female, from 9.3 km W ^a Soleil, 92 m, Departe- ment de TArtibonite, Haiti, one of a se- ries collected 20 June 1978 by Eugene D. Graham, Jr. , and Albert Schwartz. Origi- nal number ASFS V45996. Paratypes (all from De'pt. de iArtibo- nite, Haiti). ASFS V45971-95, ASFS V45997-6012, same data as holotype; ASFS V45935, Terre Sonnain, 1.6 km N Les Poteaux, 122 m, 7 August 1977, na- tive collector; ASFS V46015, 7.7 km W ^a Soleil, 30 m, 20 June 1978, E.D. Gra- ham, Jr. ; ASFS V46338-49, Lapierre, 7.6 km W ^a Soleil, 122 m, 9 July 1978, na- tive collectors; ASFS V46503-12, Lapi- erre, 7.6 km W ^a Soleil, 122 m, 10 July 1978, native collectors; ASFS V46588-91, 5.6 km W ^a Soleil, 11 July 1978, native collectors; ASFS V46603-13, 15.9 km W fa Soleil, 11 July 1978, native collector; ASFS V46615-19, 9.3 km WC^a Soleil, 11 July 1978, native collectors; ASFS V46691-93, 12.2 km W fa Soleil, 12 July 1978, native collectors; ASFS V46768-93, No. 1 Hispaniolan Sphaerodactylus 12.2 km W ^a Soleil, 14 July 1978, native collectors; ASFS V46594, ASFS V48012- 1 19, Lapierre, 7.6 km W ^a Soleil, 122 m, 11 July 1978, ASFS V46697. ASFS V48200-10, ASFS V48221-97, RT 5618- 63, AMNH 115517-31, ANSP 27170-79, CM 60522-36, KU 93393-402, MCZ 132392-400, UF/FSM 21587-600, USNM 197339-55, Lapierre, 7.6 km W ^a Soleil, 122 m, 12 July 1978, native collectors. Definition. An Hispaniolan species of Sphaerodactylus characterized by the combination of: 1) dorsal scales keeled and slightly imbricate, without a middor- sal row of small scales; 2) low number (33-42) of scales at midbody; 3) modally 2 internasal scales; 4) dorsal pattern of two dark brown dorsolateral and two dark brown lateral lines on a purplish tan ground color, the dorsolateral lines with a series of tiny white dots, each encompas- sing a granule (on the neck) or a portion of a dorsal scale (on the body), these white dots numbering as many as 12 on each side; 5) a dark brown nuchal Y, which may disappear or become more diffuse with increasing size or darker ground color, the Y most often repre- sented only by its stem in males, complete in females (Fig. 2); 6) heads yellowish to yellow in males, only faintly yellowish in females, throat and tails bright yellow in males; 7) often with some sort of stippled, hneate or reticulate dark throat pattern in both sexes; more pronounced and fre- quent in males. Description of holotype. An adult fe- male with a snout-vent length of 30 mm and a tail length (distal three-quarters re- generated) of 22 m. Scale counts are: dor- sals between axilla and groin 31, ventrals between axilla and groin 31, scales around midbody 40, fourth toe lamellae 12, supralabials to mid-eye 4/4, 2 inter- nasals. Dorsal scales subimbricate to not imbricate, keeled; ventral scales smooth, imbricate, cycloid. Dorsum in life pale tan to purplish tan with a pair of dark brown dorsolateral stripes from level of the upper margin, of the eye onto the un- regenerated base of the tail, a pair of similarly colored lateral stripes from about the level of the upper lip posteriorly above the forelimb insertion to above the hindlimb insertion, much less clear on the base of the tail, and the vague r'J B '■I i^fir^ Fig. I. Hispaniola, eastern Haiti and western Republica Dominicana, showing the known distributions of Sphaerodactylus of the shrevei group: hexagon, S. shrevei; squares, S. asterulus; circles, 5. rhabdotus; triangles, S. leiicaster. Some localities for 5. asterulus have been omitted for clarity. 10 Tulane Studies in Zoology and Botany Vol. 22 remnants of a third pair of lines low on the sides separating the dorsal tan ground color from the grayish flesh-colored ven- ter; a nuchal Y present but very diffuse, the stem abbreviated; a series of white dots in the dorsolateral lines beginning on the postocular granules (where each spot is restricted to a granule) and proceeding posteriorly, about 6 dots on each side; fore- and hindlimbs mottled with shades of brown to tan; head faintly yellowish and without distinct pattern except for dark lores and dark upper surface. Variation. Of the series of 448 S. as- terulus, the complete suite of counts has been taken on 63 specimens; counts of supralabials and intemasals, and snout- vent length measurements only were ta- ken on the remaining specimens. The largest male (RT 5628) has a snout-vent length of 31 mm, the largest females (ASFS V46776, ASFS V48123) 31 mm. Scale counts are: dorsals axilla to groin 17-24 (x = 21.4); ventrals axilla to groin 25-33 (28.6); midbody scales 33-42 (37.2); supralabials to mid-eye 4/4 (414 individuals), 3/3 (2), 3/4 (2), 4/5 (25), 5/5 (3); intemasals 2 (266 individuals), 1 (50), 3 (107), 4 (2); fourth toe lamellae 8-12 (10.2; Mq = 10 - 37%); escutcheon 1-9X6-21. ij^^,:r^''r- . ■ -' *m i. Fig. 3. The type-locality of 5. astendus (9.3 km W fa Soleil, Dept. de I'Arbonite. Haiti); from a Koda- chrome slide taken by Eugene D. Graham, Jr., 14 July 1978. In such a long series, including large numbers of males, females, subadults, and juveniles, it is not surprising that there is great variation in dorsal color and pattern. As a general statement, males are pale purplish tan dorsally, with 2 dark brown dorsolateral lines and a pair of lateral brown lines, to give a generally tetralineate effect; in addition to these lines, there is a ventrolateral pair, that are diffuse or broken, at the junction of the dorsal and ventral colors; the dorso- lateral lines have as many as 12 tiny white dots enclosed within them, these dots en- compassing a single granule on the neck region (where there may be 3 or 4 white dots crowded together) and only a por- tion of a dorsal scale on the body. These white dots are tiny but nevertheless quite conspicuous in life against the brown stripe. The cephalic Y is present but in males may be reduced to only its stem as a short middorsal nuchal dark brown line. The heads of males are yellow to yellow- ish, and the throats are bright yellow as are the tails. The venter is grayish flesh- colored, in sharp contrast to the yellow throat. Females are basically Hke the males in dorsal pattern but usually with a com- plete cephalic Y; the heads are at times only faintly yellowish. The general fe- male dorsal aspect is paler than that of males, more contrastingly lined, and the dorsal ground color tends towards tans without purplish tints. The iris in both sexes is yellow. Juveniles are patterned like females, but the ground color is paler, and the lines are all more distinct. In juvenile specimens the ventrolateral pair of lines may be as distinct and entire as the dorso- lateral and lateral lines, or they may be fragmented and incomplete as described for the holotype. The dorsolateral lines are common to most of the series. In some females, the No. 1 Hispan iolan Sphaerodactyhis 11 contrast between the lines and the ground color is not so intense as in other females, and the occipital Y is either incomplete (represented by its stem only) or diffuse and almost concolor with the dorsal ce- phalic brownish color. Throats in both sexes may be immaculate. This is the common condition in males, whereas in many (but not all) females there may be vague indications of a grayish reticulum or grayish stippling on the throat. In sub- adult females the dark loreal line and a dark median snout line are present, but these are obscured with increased size. There is little tendency for the dorsolat- eral lines to break up into a series of blotches (as in S. shrevei). and the lines are almost always entire. Occasion- al specimens show a tendency toward fragmentation but these are rare. If so, the dorsolateral line fragments are not blotchlike and are recognizable as parts of dorsolateral lines, not independent blotches. The most distinctive feature of S. as- terulus is also the most variable in pre- served specimens: the tiny white isolated dots in the dorsolateral lines. In our origi- nal series of 42 specimens, our color notes state that all specimens (23 males with snout-vent lengths between 22 and 29 mm; 18 females with snout-vent lengths between 21 and 30 mm; one hatchling with a snout-vent length of 15 mm) had dots present in the dorsolateral lines. When these (and the remainder of the long series) were studied approximately 4 months later, most of both sexes still showed the white dots or the scales where they had been, but they were less often encountered in males (53%) than in fe- males (86%). Most persistent are the dots on the granular neck scales, with those on the body tending to become obscure or lost. Since preserved S. leucaster com- pletely lose their white "stars" promptly after preservative, we can logically as- sume that the same is true for S. asterulus and its dots. A second factor is that the dots in S. asterulus tend to be less clear and bold in large specimens. Both these factors, then, would tend to account for the loss of white dots in such a large per- centage of S. asterulus. (It is interesting that juveniles with snout-vent lengths of 19 mm or less retain spots clearly.) But these factors do not account for the fact that almost half of the male 5. asterulus (47%) have lost their dots, whereas a much smaller percentage (14%) of the females has done so. We suggest that there is some weak dichromatism in 5. asterulus with more males than females tending toward an unspotted condition in life, and that by chance our original series had all males with dots. Comparisons. Sphaero dactyl us asteru- lus is geographically close to S. shrevei (80 km) across the Presqu'ile du Nord Quest and is patterned similarly to S. leu- caster, from whose range S. asterulus is removed from 220 km. From 5. shrevei, S. asterulus differs in having a lined dor- sum in contrast to a blotched or cross- banded one. Sphaerodactylus shrevei av- erages higher in all scale counts (dorsals 23.8 versus 21.4 in 5. asterulus; ventrals 32.2 versus 28.6 in S. asterulus; midbody scales 42.0 versus 37.2 in S. asterulus), although all counts overlap broadly with least overlap in number of dorsals. The modally 1 intemasal in 5. shrevei differs from the modally 2 intemasals in S. as- terulus (71% versus 60%); 50 of 445 (11%) 5. asterulus have 1 internasal. 5. asterulus and S. leucaster are in some ways similar - both have dorsolateral lines with white inclusions - dots in the former case, large radiating white spots in the latter. In S. leucaster, the white spots interrupt the dorsolateral lines complete- ly, resulting in irregularly broken dorso- lateral lines, whereas this is not the case in S. asterulus. S. asterulus also has well 12 Tulane Studies in Zoology and Botany Vol. 22 developed lateral stripes, which are ab- sent in S. leucaster. Scale counts and fre- quencies are comparable. From 5. rhabdotiis, S. asterulus differs in having dots in the dorsolateral lines, whereas S. rhabdotus has neither spots or dots - the lines are entire (see, however, the discussion of 5. rhabdotus variation in the present paper). S. asterulus has a lower mean in dorsal scales (21.4) and midbody scales (37.2) than either 5. leucaster (23.5 and 39.8) or S. rhabdotus (23.8 and 40.4). On the oth- er hand, S. asterulus has a higher ventral mean (28.6) than either S. leucaster {21 A) or S. rhabdotus (28.0). In all cases the ranges of variation overlap broadly. Two internasals show about the same fre- quency in all three species (5. asterulus 60%, S. leucaster 68%, S. rhabdotus 63%). Both S. asterulus and S. leucaster have modes of 10 subdigital lamellae, whereas S. rhabdotus has a mode of 9. The modes are weak (37% and 44% in the first two species, 36% in S. rhabdo- tus). Means of fourth toe lamellae vary from 9.5 in S. leucaster to 10.2 in 5. aster- ulus. Escutcheon size in males of all three species is peculiar in its extreme variabil- ity. In S. asterulus^ the length varies from 1 to 9 scales, and the width from 6 to 21 scales. The length of the escutcheon in male 5. leucaster (5-8) and 5. rhabdotus (5-9) is much less peculiar, but in these two species the width of the escutcheon shows extreme variability, from 8-24 in 5. leucaster and 7-25 in S. rhabdotus. On- togeny and sexual maturity may be fac- tors in this case. Members of the shrevei complex have only hair-bearing scale organs. In S. as- terulus the number is extremely variable, with between 6 and 15 organs along the free margin of the scale, each with from 1 (usually) to 2 or 3 "hairs." S. rhabdotus has 5 to 1 1 hair-bearing organs, each with 1 "hair," per scale (Schwartz, 1970:64) and S. leucaster has 3 to 8 organs, usually \ with 2 "hairs'' (Schwartz 1973:36). Remarks. We have already mentioned the taking of the original series of S. as- terulus within the dead basal rosettes of Agave and in low rock piles. In the debris in the Agave leaves we also encountered many eggs, mostly hatched, but five un- hatched. Measurements of four of these vary (in mm) between 6.4-6.8 X 5.1-5.7. The fifth egg hatched in the collecting bag, and the juvenile from it has a snout- vent length of 15 mm. In addition to the large numbers of S. asterulus encoun- tered by us, we also secured an adult Epi- crates f. fordi Gunther and an immature Phyllodactylus. The snake was within the punky stem of an Agave, whereas the lizard was in a pile of rocks and Agave leaves. DISCUSSION The shrevei group is composed of four species whose range occupies two general regions: extreme northwestern Haiti and the Valle de Neiba and the Llanos de Azua in the Republica Dominicana. Those authors who have previously dealt with members of the group (Lazell, 1961 ; Schwartz, 1970, 1973) have been hard pressed to suggest to what other Hispani- olan or even Antillean geckos the shrevei group is related. The combination of characteristics of the shrevei group set it off from all other Sphaerodactylus; the convex snout and vertically elliptical pu- pils are, in combination, unique. These characters are apparently primitive with- in Sphaerodactylus, and it may well be that the shrevei group represents an old remnant of a Sphaerodactylus radiation. Its apparently fragmented distribution would tend to confirm this interpretation. It is also possible that the shrevei group is derived from the difficilis complex, which No. 1 Hispaniolan Sphaerodactylus 13 is in turn composed of the Hispaniolan members of the notatus group. We make this latter suggestion, not for any particu- lar reasons of similarity between the two (although at least keeled dorsal scales is one character in common between them), but only because the difficilis complex is the most widespread and most diverse of the Hispaniolan groups and/or complexes of geckos. Although the dif- ficilis complex lizards do not totally shun xeric regions, they are not the "typical'' geckos occupying xeric areas. Thus the shrevei group and the difficilis complex are roughly complementary in distribu- tions. One member of the difficilis com- plex. S. cdtavelensis Noble and Hassler, is a xerophile, and it has been taken sym- patrically with S. asterulus; another member, S. cryphius Thomas and Schwartz, was found in the same thatch pile as the type series of S. rhabdotiis. Sphaerodactylus ocoae Schwartz and Thomas also appears to be at least margi- nally sympatric with S. leucaster, and S. difficilis Barbour and S. leucaster may well be sympatric in portions of the Lla- nos de Azua. A still-unnamed species of Sphaerodactylus, not of the difficilis com- plex, has been taken with S. asterulus. Despite the above examples of macro- sympatry and even syntopy, but on the other hand taking into consideration the large number of Hispaniolan Sphaero- dactylus species, we can say as a general statement that members of the shrevei group are so confirmedly xerophiles that they are rarely found with other Hispani- olan Sphaerodactylus. It would be most convenient to state that the two northern species {S. shrevei and S. asterulus) were more closely re- lated to each other than they are to the southern pair {S. rhabdotus and S. leucas- ter: see Fig. I ). This seems not to be the case; the lineate pattern of 5. asterulus, S. rhabdotus, and S. leucaster wou\d seem to ally them to each other, despite the wide hiatus between them, rather than the blotched pattern of S. shrevei and the dotted pattern of S. asterulus. [iven more convincing of this relationship are the white dots of S. asterulus and the white blotches ofS. leucaster. In fact, one might with some justification consider 5. asteru- lus a subspecies of S. leucaster on this basis. But the wide geographical separa- tion of these two taxa, plus the interposi- tion between them of 5. rhabdotus (which lacks white markings in the dorsolateral lines) suggests that we are dealing with three species. We have commented on the complementarity of the ranges of S. leucaster and S. rhabdotus and the possi- bility that perhaps these two taxa might better be regarded as subspecies; certain- ly the 21 km gap between their known ranges is small. But between them cours- es the large Rio Yaque del Sur, a major river, and S. leucaster occupies areas east of the river and S. rhabdotus the west. Williams (1961) noted that the Hispan- iolan herpetofauna is divided into two major sections, corresponding to the north and south paleoislands into which modern Hispaniola was divided. This point has been emphasized repeatedly by other authors, and it is a valid one. Al- though there are exceptions, most of the Hispaniolan herpetofauna can be divided into north and south island elements; the modern division of these two old islands is the Cul de Sac-Valle de Neiba plain, and it is in the latter portion of this (the Valle de Neiba) that S. rhabdotus occurs. There is no question that the shrevei group as a whole is north island in af- finities: three of its species (-S". aster- ulus. S. shrevei. S. leucaster) occur exclusively upon it. But S. rhabdotus, originally described from a south island locality, has now been taken on both sides of the Valle de Neiba; two localities (Las Lajas; La Descubierta) are on the 14 Tulane Studies in Zoology and Botany Vol. 22 northern side, whereas three others (La Florida; Duverge; Angostura) are on the south. If we postulate that S. rhabdotus and S. leucaster were originally inhabit- ants of the southern xeric shore of the north island, the two species separated by the north island Rio Yaque del Sur, then it is probable that, with closure of the interisland strait, S. rhabdotus has cros- sed the Valle de Neiba to reach to the foothills of the Sierra de Baoruco on the south island. Such a scenario presents no major problems and accounts for the oc- currence of S. rhabdotus away from the north island center of the group. S. shrevei and S. asterulus are appar- ently restricted to northwestern Haitian Presqu'ile du Nord Quest, the former on the peninsula itself, the latter at its south- eastern base. Most of this peninsula re- mains unknown herpteologically, and we have as yet no clear concept of what spe- cies occur there or the precise distribu- tions of the species already known. Most of the latter are known from very limited material and from a few scattered locali- ties (or only one locality). What is puzzl- ing are the radical differences in pattern between S. shrevei and S. asterulus — blotched in the former, lineate in the lat- ter. Thus, S. asterulus much more closely resembles in pattern the more distant S. rhabdotus-S. leucaster pair than it does adjacent S. shrevei. This may only mean that 5. shrevei has been isolated for a much longer period from the basal, Une- ate shrevei group stock than has 5. as- terulus. Until more material of both spe- cies is known from this region, it is futile to speculate further. We should also like to point out that the road between the main road near Les Poteaux and the northern coast at Port-de-Paix passes through xeric country for much of its 77 km; only toward its northern coastal sec- tion near Port-de-Paix may the area be called mesic. Yet we had no success in securing shrevei-group geckos along this road. We did encounter S. difficilis at several xeric locahties, which suggests that no shrevei-group member occurs here. But since shrevei group sphaero- dactyls occur in very rigorous situations and may not be common, our not secur- ing any in this region may not be so signi- ficant. The 130 km hiatus between S. asterulus and S. rhabdotus may be more apparent than real. Sphaerodactylus rhabdotus is still known from the Haitian Cul de Sac plain, although it occurs very near the border (Las Lajas). We confidently ex- pect it in this valley. From the western end of the Cul de Sac, xeric coastal areas follow the shore of the Golfe de la Go- nave to near St. Marc. Just north of that city lies the wide (30 km) Vallee de I'Ar- tibonite, presently irrigated and flooded for the cultivation of rice. Inland, how- ever, the Vallee de I'Artibonite is xeric and could supply suitable habitat for a shrevei group member. North of the val- ley and southeast of the city of GonaiVes, there are once more extensive xeric areas which are continuous with the region where S. asterulus occurs. It would seem likely that 5. rhabdotus occurs from the Republica Dominicana into the Cul de Sac plain and thence along the shore to the southern edge of the Vallee de I'Arti- bonite and that S. asterulus occurs as far south as the northern edge of this same valley. But efforts on our part and those of others to secure members of this group in this region have failed totally. In fact, the sphaerodactyl fauna of this region is poorly known. As in all desert regions, native cooperation is hard to encourage, and collecting on the part of the investi- gator is more than likely to reveal nothing for lack of suitable places to search. Cer- tainly this entire region is one which de- serves special attention as far as its Sphaerodactylus fauna is concerned. No. 1 HispaniolanSphaerodactylus 15 ACKNOWLEDGMENTS In addition to specimens in the Albert Schwartz Field Series (ASFS), we have examined material in the Carnegie Mu- seum (CM), Museum of Comparative Zoology (MCZ), Texas Cooperative Wildlife Collection, Texas A and M Uni- versity (TCWC), University of Florida, Florida State Museum (UF/FSM), and the National Museum of Natural History (USNM). We wish to thank the respec- tive curators and their assistants for loans of material: Clarence J. McCoy, Ernest E. Williams, Jose P. Rosado, James R. Dixon, Walter A. Auffenberg, Peter Meylan, George R. Zug, and Ronald I. Crombie. In the field we have had the assistance of Danny C. Fowler, Mark D. Lavrich, John C. Lucio, James A. Rod- gers, Jr., Bruce R. Sheplan, William W. Sommer, Michael H. Strahm, James B. Strong, and Richard Thomas. Father John Breslin made Lucio's and Sommer's stay at Mole St. Nicholas pleasant and was instrumental in their securing the fresh material of S. shrevei. Paratypes have been placed in some of the above collections and the American Museum of Natural History (AMNH), the Academy of Natural Science of Philadelphia (ANSP), and the Museum of Natural History, University of Kansas (KU). We have also examined specimens in the col- lection of Richard Thomas (RT). Some specimens of S. rhabdotus and S. leucas- ter were collected under the auspices of National Science Foundation grants G- 7977 and B-023603 to the senior author. We are especially grateful to Fred G. Thompson and Ronald I. Crombie for keeping us abreast of their activities in the Repiiblica Dominicana and for allow- ing the senior author to examine material promptly after it had been collected. The illustrations of the four shrevei group spe- cies are the work of Beryl Bayer to whom we are grateful. LITERATURE CITED Lazell, J.D. 1961. A new species oiSphaerodactylus from northern Haiti. Breviora, Mus. Comp. Zool. (139): 1-5. Schwartz, A. 1970. A new species of gecko (Gek- konidae, Sphaerodactylus) from Hispaniola. J. Herpetology, 4: 63-67. . 1973. A third species of the Hispani- olan shrevei group of Sphaerodactylus (Sauria, Gekkonidae). Proc. Biol. Soc. Washington, 36: 35-40. Williams, E.E. 1961. Notes on Hispaniolan herpe- tology. 3. The evolution and relationships of the AnoUs semUineatus group. Breviora, Mus. Comp. Zool. (136): 1-8. 16 Tulane Studies in Zoology and Botany Vol. 22 NATURAL HISTORY OFTHE MAP TURTLES CRAPTEMYSPSEUDOGE(X,RAPHKA AND T;. OUACHITENSIS IN WISCONSIN RICHARD C. VOGT ih\ivcrsii\- o'f Wisconsiii-Mculisoii. '/.oology Dcpartiucnt^ Madison. Wisconsin ABSTRACT Natural history data were collected from popula- tions of Graplemys ouachitensis and C psemlogeo- graphica on the Mississippi River, Vernon County, Wisconsin, from 1972-1978. Clutches of eggs were incubated in the laboratory and field under various temperature regimens. Hatchlings were maintained in the laboratory for six years to study growth rates, size at reproductive maturity and sex ratio within clutches. Examination of reproductive tracts of 50 females suggest that they lay two clutches of eggs annually. The average clutch size for G. pseudogeographica was 14.1 and for G. ouachitensis, 19.2. The court- ship displays of male G. ouachiiensis and G. pseu- dogeographica involve the drumming of the fore- claws against the ocular regions of the female. The two species differed in the number of contacts per "titillation" bout. Nesting was observed from 18 May to 26 July. Nest temperatures were monitored continuously for one month in 1972. Incubation temperatures in the laboratory were shown to affect hatching success, number of scute anomalies, size of vellow head blotches, and sex determination in both species. A skewed sex ratio of five females per male was cal- culated from trapping results. Stomach contents of adult females suggested that the food resources are being partitioned. Females do not begin feeding until after laying their first clutch of eggs for the season. The primary function of basking is thermoregu- latory, no aggressive interactions were observed be- tween basking Graptemys. Crackles, Quiscalus quiscala. were observed removing leeches, Placob- Present Address: Carnegie Museum of Natural History, Section of Amphibians and Reptiles. Pitts- burgh, Pennsvlvania 15213 della parasitica, from basking Graptemys. Nest pre- dation by red fox, raccoon, and otter was observed. INTRODUCTION The false map turtle complex. Grapt- emys p. pseudogeographica, G. o. ou- achitensis, G. o. sabinensis and G. p. koh- ni has perplexed taxonomists and con- fused ecologists since the time of the orig- inal descriptions ( Vogt, in press). Ecologi- cal data involving Graptemys have often been based on populations that were in- accurately identified. I realized this prob- lem while attempting to determine which species of this group occurred in Wiscon- sin. Specimens collected from the Missis- sippi River near Stoddard, Vernon Coun- ty, indicated that all taxa listed above occur sympatrically in Wisconsin. In 1971 at the same locality I collected 26 clutches of eggs from unknown females. Labora- tory incubation yielded hatchlings with morphological characteristics of each, suggesting that pseudogeographica, koh- ni, ouachitensis and sabinensis have been described from individuals of a single highly polymorphic species. Smith (1961: 150-151) suggested this as one hypothe- sis, having been confronted with the same problem with false map turtles in Illinois. These species and subspecies have been defined primarily on differences in head markings (e.g. Cagle, 1953; Ernst and Barbour, 1972); using these criteria EDITORIAL COMMITTEE FOR THIS PAPER: DR. CARL H. ERNST, Professor of Biology, George Mason University, Fairfax, Virginia 22030 DR. EDWARD O. MOLL, Professor of Zoology, Eastern Illinois University, Charleston, Illinois 61920 17 18 Tulane Studies in Zoology and Botany Vol. 22 all "forms" were found to have emerged from the 26 clutches of eggs, in some cases more than one "species" or "sub- species" from a single clutch. Study of this taxonomic perplexity was begun in 1972 at the Stoddard site where the population of Graptemys is large, and accessible. Be- tween June and August of that year a mark and recapture study was begun. Subsequent trips were made intermit- tently between April and November of 1972-1978. Although I was concerned primarily with systematics of the G. pseudogeographica complex, various as- pects of the natural history of Graptemys were observed. The field study was com- plemented by a five year laboratory study wherein eggs were incubated and young raised. Courtship behavior, reproductive potential and growth rates were studied in both the field and laboratory. This pa- per deals primarily with the natural his- tory of G. ouachitensis and G. pseudo- geographica. Detailed analysis of the sys- tematics, courtship behavior and re- source partitioning are presented else- where (Vogt, 1978). SYSTEMATIC RELATIONSHIPS AND DISTRIBUTION Graptemys pseudogeographica was de- scribed by Gray (1831) from the Wabash River at New Harmony, Indiana. Grapt- emys pseudogeographica is found pri- marily in large stream systems of the Mis- sissippi drainage and occurs from the St. Croix and Wisconsin rivers in northern and central Wisconsin and the upper Mis- sissippi River in Minnesota southward through Louisiana and eastern Texas (Vogt, Bull. Carnegie Museum, in press). Cagle (1953) described two new sub- species of Graptemys pseudogeographica from the southern United States: ouachi- tensis from the Ouachita River of Louisi- ana and sabinensis from the Sabine River on the Louisiana-Texas border. My stud- ies (Vogt, 1978) show that these two sub- \, species constitute a separate species, G. ouachitensis, which occurs from the Mis- sisisppi and St. Croix rivers in Wisconsin and Minnesota, south through the Missis- sippi River basin into Louisiana, west to Lake Texoma, Oklahoma and east into Indiana and West Virginia. THE STUDY AREA My study area, referred to as Missis- sippi River Pool 8, extends 37.5 river km !| north of a dam at Genoa, Vernon Coun- ty, Wisconsin, to a point north of La- Crosse (43^40' N latitude, 93^13' W Ion- * gitude). The Root and Lawrence rivers are the main tributaries of this impound- ment. Before construction of the lock and dam in 1930, the main river channel com- , prised the bulk of the water surface area: 12,597 ha of channel, 2,764 ha of slough, 4,606 ha of marsh and 1,383 ha of pond. Since the dam was closed in 1937, water in the channel rose 3.5 m creating 27,481 ha of open pool and 2,873 ha of feeder channels; the marshland was increased to 9,435 ha (Claflin, 1973). Much of the pool area was taken from what previously had been river bottom forest (Curtis, 1959), stumps of which still remain provide ex- cellent basking site for the turtles and attachment sites for invertebrates. Tim- ber-meadow in the study area was 56,810 ha in 1930, but in 1970 only 25.396 ha remained (Claflin, 1973). Within this general area, study was concentrated on 230 ha within the per- imeter defined by the intersection of Crosby Slough and the main channel to the north, Crosby Slough and Cook Slough to the south and the main channel to the west. Most nesting observations were made on two islands which border the main channel (Fig. 1). The islands are characterized by ex- panses of open sand surrounded by trees No. 1 Wisconsin Map Turtles 19 [Populus deltoides Marsh., Fraxinus pennsylvanica var. subintegerrima (Vahl) Fern., Acer negundo L., A. saccarinum L., Betula nigra L. and Ulmus americana L.] and shrubs [Salix sp., Amorpha friiti- cosa L., Parthenocissus inserta (Kerner) K. Fritsch, Vitis riparia Michx. and Cor- niis obliqua Raf.]. The trees and shrubs are essential in maintaining the integrity of the island by limiting wind and water erosion. Open sand is sparsely covered with a variety of herbs, the most common in the open sand being Sporobolus cryptandrus (Torr.) Gray, Carex sp., Bromiis tector- um L., Tradescantia ohiensis Raf., Ver- basciim thapsiis L. , Oenothera biennis L., and Lepidiiim densifloriim Schrad. Grap- temys frequently dig nests adjacent to th- ese plants. Along the border of the sand and edge of the woods Asclepias syriaca L., Barbarea vulgaris R. Br., Saponaria officinalis L., Mollugo verticillata L. and Cycloloma atriplicifoliiim (Spreng.) Coult. were common. Some plants occur primarily in the woods but were found at the open sand-woods interface: Rhus ra- dicans L., Laportea canadensis (L.) Wedd., Urtica dioica L., Solanum dul- camera L., Lynchnis alba Mill., Polygo- num convolvulus L., Erigeron annuus (L.) Pers., Equisetum arvense L. and Galium obtusum Bigel. Phalaris arundi- nacea L. occurs along the edge of the water. The shrubs and trees along the pe- riphery of the islands as well as the her- baceous plants on the open sand help pre- vent the wind from eroding the sand and exposing nests, and provide cover for hatchling turtles as they move to the water. The turtles feed on and in the common marsh aquatic plants found in shallow wa- ter. Near the nesting islands these are Scirpus validus Vahl, 5. fluviadlis (Torr.), Sagittaria latifolia Willd., Ziza- nia aquatica L., Phalaris arundinacea L., Phragmites communis Trin., Sparganium americanum Nutt. and Carex sp. In deep- er water near the islands and in the feed- ing areas near the stumps Potamogeton nodosus Poir., P. crispus L., P. ameri- canus C. & S., P. foliosus Far., Heteran- thera dubia (Jacqu.) MacM., Vallisneria americana Michx., Elodea sp., Nym- phaea tuberosa Paine, Nelumbo lutea (Willd.), Spirodela polyrhiza (L.), Lem- MISSISSIPPI RIVER - POOL 8 LA CROSSE Figure 1 . Topographic map of study area showing location of trap sites. 20 Tulane Studies in Zoology and Botany Vol. 22 na minor L., Wolffia columhiana Karst., Ceratophvllum sp. and Myriophyllum sp. provide both food and shelter. MATERIALS AND METHODS Turtles were collected in fyke nets with leads, gill nets, and trammel nets and by hand capture (Vogt, Copeia, in press). Fyke nets varying in diameter from 61 cm to 90 cm and in mesh size from 2.54 cm to 5.08 cm were set in 1-1.5 m of water with the leads parallel to the shorelines of nesting beaches, between feeding grounds and deep water, or between basking logs and deep water. Graptemys was not attracted to bait, therefore fyke nets were baited with fresh dead carp on- ly when turtles other than Graptemys were desired. Local commercial fishermen, when setting gill nets in the study area, saved Graptemys that became entangled in the 12.5-17.5 cm mesh gill nets. Many of these were freshly drowned specimens from which eggs could be removed and incubated. These dead specimens were also used for stomach content analysis and skeletal preparation. Trammel nets (5.08 cm mesh, 17.8 cm outer walling) were used primarily near basking logs or wing dams during hiber- nation periods (October, November, Ap- ril). Turtles were driven into the nets by the use of a "carphorn" (Vogt, 1978). Female turtles were hand captured on nesting beaches and by snorkeling while they were feeding. Hatchlings were col- lected in shallow water by hand and dip- net. Seining with a 60 m, 2.54 cm mesh bag seine proved to be ineffectual for ob- taining young or adults. During the first summer of studv (1972) clutches of eggs from known fe- males of G. ouachitensis and G. pseiido- geographica were collected for taxonom- ic study. All six nesting beaches on the island were monitored each morning from 0500-0830 hrs (Central Standard Time) to obtain females and their recent- ly laid egg clutches. Eggs from known females were measured by use of dial cal- ipers accurate to 0. 1 mm and incubated in the laboratory at 280C. Each clutch was labeled and kept in a separate glass bowl containing moist spaghnum moss; and the entire bowl was enclosed in a clear plastic bag to prevent desiccation. Following morning surveillance of nesting beaches, the 12 to 18 fyke nets, set at various localities within the study area, were checked (Fig. 1). All G. psen- dogeographica and G. ouachitensis were brought back to camp for marking, and recording of data. Each turtle was weighed, measured, photographed and marked; notes were made on shell injuries, abnormalities, color patterns and location of capture. Females were palpated for the presence of shelled eggs. Weight was determined to the nearest gram. The turtles were weighed under field conditions; upon handling Graptemys usually void their bladder. Injection of pilocarpin (Doran- do, 1979) to induce voiding of all bladder water was not feasible or desirable since the inherent error was shown to be only 2.43% and any lasting effects on physiol- ogy or behavior of the turtle have not been studied. Maximum straight line car- apace length, width and height were mea- sured to the mm by use of a turtle measur- ing board (Cagle, 1946). Plastral mea- surements were taken either with dial cal- ipers accurate to 0. 1 mm or with a flexible ruler (nearest mm). Plastron width was taken at the suture between the axillary and pectoral scutes where it meets the 5th and 6th marginals. Straight line plastron length was taken from the farthest exten- sion of the gular and anal scutes. During the summer of 1972 only carapace length and width were recorded. Both sides, dorsum, and venter of the head of each No. 1 Wisconsin Map Turtles 21 turtle were photographed. Each turtle was individually marked with colored plastic plugs inserted with a "Dennison Buttoneer" through a 2 mm hole drilled through the center of one of the 7th-12th marginals on each side of the carapace. The marked turtles were then released at the site of capture the following day. Fe- males were sacrified from April-Novem- ber to obtain eggs and fresh reproductive tracts for determining reproductive con- dition. Notes were made concerning the number of growth rings on an abdominal scute when discernible. A ring was mea- sured from the edge of the areola to the first deep groove in the scute. In agree- ment with Moll and Legler (1971) the term annulus is not used since it implies yearly rings and no proof exists to show if rings actually represent only one year. Eggs were obtained by induction of oviposition by oxytocin injections (Ewert and Legler, 1978) during May and June of 1976 and 1977. If shelled eggs could be discerned by palpation, the females were taken back to the laboratory and injected intramuscularly with 10 units of oxytocin, then placed in a container of clean water, 25 cm deep at 28°-32°C and kept under bright lights with minimal disturbance. If eggs were not laid within one hr an addi- tional injection of 10 units was given. Weights were taken before and after egg laying. The eggs were then weighed, measured and placed in moist vermiculite in incubators set at 25°, 30°, or 350C or placed in a laboratory room where tem- perature was maintained at 32°C with a 14 hr light cycle. In 1977, individual eggs of seventeen clutches were incubated at these different temperatures. Young from the 1972 year class have been reared in the laboratory to October 1978 (48 surviving) on a prepared diet of agar, gelatin, trout pellets, bone meal, oyster shells, cod liver oil, and multiple vitamins. This was done to compare growth rates between males and females and to determine if any changes in head color patterns occurred. They have been kept on a 14 hour light cycle at 32°. "Vit- alities" (Durotest) were used to provide ultraviolet light. Blocksof plaster of Paris and copper tubing were kept in each tank to assist shell development. All the young hatched in 1972 were measured, weighed, marked and photographed in a manner similar to that used with adults. In addition, photographs were taken of the carapace and plastron. Measure- ments were repeated at least once a year. After the second year of growth, when external sex characteristics became appa- rent, males and females were placed in separate containers. Analysis of feeding preference was made by examination of the stomach con- tents of freshly sacrificed or drowned in- dividuals. Stomach flushing (Legler, 1977) was used during the 1977 season. Stomach contents from females were sorted and the components identified. Volume of the sorted contents was then determined by water displacement (Vogt, 1978). Feeding behavior of fe- males was observed through a 30X spot- ting scope, or by snorkeling in their feed- ing areas. Nest and body temperatures were ta- ken with a Schultheis quick reading ther- mometer. An Easterline-Angus six- channel recorder with six thermistors was used for continuous recording of the temperature in four nests and the sand and air surface from July 16, 1972 through August 17, 1972. All turtles sacrificed or drowned in the nets of commercial fishermen were either fixed in 109"^ formalin or skeletonized by dermestid beetles and deposited in the University of Wisconsin Zoological Mu- seum, Madison. Hatchlings from 1971, 1972, 1976 and 1977 were also preserved and deposited there. 99 Tulane Studies in Zoology and Botany Vol. 22 Table 1 = Measurements (cm) of carapace length (CL), carapace width (CW), cara- pace height (CH), and weight (g) of male and female Graptemys from Stoddard. (Range above; mean below) Species Sex n CL CW CH WT (g) G. ouachitensis G. geographica G. ouachitensis d G. psueodgeographica d d G. pseudogeographica + G. geographica 68 10.9-13.7 8.7-10.3 4.0-5.2 161-309 (12.3) (9.3) (4.5) (211) 68 11.1-15.1 8.7-11.0 4.0-5.3 143-364 (13.3) (10.0) (4.7) (251) 45 9.3-13.6 7.6-11.1 3.4-4.6 165-350 (11.5) (8.9) (3.9) (205) 265 16.3-24.2 12.7-19.1 6.5-10.2 557-2300 (20.5) (15.8) (8.5) (1136) 109 19.3-27.4 15.1-20.4 7.3-11.2 857-2502 (22.5) (17.3) (8.8) (1477) 15 20.1-25.8 14.9-18.7 7.5-9.4 761-1700 (22.6) (16.9) (8.7) (1138) REPRODUCTION Sexual dimorphism Sexual dimorphism is pronounced in all species of Graptemys. Males are much smaller in carapace length, width and height (Table 1). The male tail is com- paratively much longer than that of the female, the distance from plastron to clo- aca being greater in the male to facilitate intromission of the penis during copula- tion. The 2nd and 3rd foreclaws of male G. ouachitensis and G. pseudogeograph- ica are disproportionately elongated. This characteristic has probably evolved along with their characteristic courtship display wherein the male drums his fore- claws over the ocular and otic regions of the female. The foreclaws of G. geo- graphica males are not elongated and their courtship display does not involve foreclaw drumming. Such a marked sexual dimorphism (males averaging 20% the mass of fe- males) has many advantages. Most im- portantly it allows for intraspecific parti- tioning of food resources. Males feed on small, energy-rich insect larvae and mol- lusks, while the adult females are omni- vores that feed on invertebrates and veg- etation. Small size may allow males to put their energy into searching for females, precopulatory display and sperm produc- tion, rather than into growth. Laboratory studies show that reproductive maturity is reached in males at a smaller size allow- ing for a lower minimum age of reproduc- tion (see section on laboratory growth, below). Females, on the other hand, have an advantage in being large. The larger the female the greater the number of eggs she produces in a single clutch (Figs. 2 and 3). Warm spring temperatures allow ovi- ducal application of the shell of the first clutch of eggs starting the last 10 days of May; then 10 days to 3 weeks to apply shell material to the second clutch, and an additional 10 days to three weeks for a possible third clutch. This is the max- imum number of clutches since tempera- ture becomes too low for incubation to be completed by the onset of winter. A larg- No. 1 Wisconsin Map Turtles 23 er female provides a higher than average reproductive input into the population. In addition the larger size of the females may afford greater protection from avian and mammalian predators when they venture ashore to nest. Reproductive cycles Females begin the reproductive cycle from late June to mid-July after laying their last clutch of eggs for the year. From then until early September females feed and bask, converting the energy obtained into enlarged ovarian follicles. They en- ter hibernation with the coelomic cavity encroached upon by enlarged ovarian fol- licles. They do not feed in the spring until the first clutch has been laid. Mating probably takes place either in October or April while the turtles are congregated at 6 pseudogeographica 18 G ouachi tensis o 14 (- _l O o) 12 UJ Ll_ O 10 cn UJ CD i 8 6- n = 65 • • mm • • • • • •• • • • mm • mm • • • m •• •• 16 18 20 22 24 CARAPACE LENGTH (cm) Figure 2. Relationship between carapace length of female and number of eggs per clutch in G. oiai- chilensis. 22 20 X o u o 18 16 n = 50 • • O 14- q: UJ m 33 12- 10- 8 •••• • • ••• • • • • 1 1 1 • 1 1 • 1 1 1 1 18 20 22 24 CARAPACE LENGTH (cm) 26 Figure 3. Relationship between carapace length of female and number of eggs per clutch in G. pseudogeographica. hibernacula. After leaving the hibernac- ula in April the females move toward the nesting beaches and spend approximately six weeks basking and applying shell to ovarian follicles, and egg laying continues until late June to mid-July. This appears to be the typical pattern for North Ameri- can emydines (Risley, 1938; Atland, 1951; Miller, 1959; Moll and Legler, 1971; Christiansen and Moll, 1973; Moll, 1973; Shealy, 1976). Testis size was noted to be greatest in the fall and smaller in the summer. Shealy (1976) found active sperm present in the epididymis of ma- ture male G. pulchra throughout the year, suggesting that mating could occur throughout the year in Alabama. Mating is potential in Wisconsin in both fall and spring. Males of all three species cap- 24 Tulane Studies in Zoology and Botany Vol. 22 tured in October, November and April courted and attempted copulation when returned to an outside enclosure at the laboratory. Courtship by laboratory raised males and males caught in the wild was ob- served in the laboratory (Vogt, 1978), and high speed motion pictures were made to analyze differences among the three species. Males identify conspecific females by both visual and olfactory cues. Specific odors appear to be associated with the cloacal region of females since a male, after placing his nostrils in close proximity of the cloaca of a conspecific females, often swims rapidly to her head and begins courting; however, evidence for long distance pheromones has not been substantiated (Vogt, 1979). Courtship in G. ouachitensis and G. pseudogeographica also seems to be in- itiated by visual cues. Males have been observed to court conspecific females and often other males after orienting towards the head without initial cloacal contact. Once a turtle is suspected to be a con- specific female, males may immediately try to mount or they may begin courtship. The courtship displays of the three spe- cies differ significantly and are probably responsible for maintaining genetic isola- tion between species pairs. The aquatic displays of G. ouachitensis and G. pseudogeographica are remarkab- ly similar. The male approaches the fe- male anteriorly with his forelimbs raised above his head and arched laterally. Only the hindlimbs are used to bring him into position. When nose to nose contact is nearly made, (within 3 mm) the forelimbs are brought down, the venters of the fore- feet are rotated laterally and the fore- claws are simultaneously drummed against the ocular region of the female head. The duration of the drumming does not differ significantly between the two species but the frequency does. Jackson and Davis (1972) described this as the "titillation" phase of the courtship be- havior in Pseudemys scripta. The length of the titillation phase was highly variable in both species of Graptemys ranging from 280-750 ms (mean 454) in 23 G. ouachitensis and 344-834 ms (mean 470) in 13 G. pseudogeographica. The mean number of contacts per bout, however, was different for the two species. The mean number of contacts for G. pseudo- geographica, 10.3, was nearly twice that of G. ouachitensis, 5.2. In addition G. pseudogeographica dif- fers by bobbing its head up and down while G. ouachitensis holds its head sta- tionary and vibrates the foreclaws. Mounting is attempted whenever the fe- male is almost motionless and does not seem to be dependent on the number of courtship bouts. Courtship in G. geographica differs in that no vibration of the forelimbs has ever been observed. The male may attempt to mount after cloacal contact. Or, after cloacal contact, the male has been ob- served swimming to the head of the fe- male and, after making snout to snout contact, bobbing his head rapidly in the vertical plane. This is similar to the court- ship behavior of G. pulchra (Shealy, 1976). Mounting is facilitated in all three spe- cies by the male hooking the tip of his tail around that of the female to maintain balance and to pull her cloaca into posi- tion to allow intromission. Once intro- mission is achieved the pairs have been observed in coitus from 15 sec to over four hrs. During this time the male re- mains in a passive state floating at ap- proximately a 45° angle upon the cara- pace of the female. His forelimbs are usu- ally arched above his head and held mo- tionless. Reproductive potential Detailed reproductive data were only No. 1 Wisconsin Map Turtles 25 Table2. Relationship of both carapace length (CL)andaveragenumbcr of eggs perclutch(n = # of clutches measured) and how they relate to size and weight of eggs in Graptemvs pseudogeographicii. Ave. no. Ave. egg Ave. no. Ave. egg Ave. no. Ave. clutch Ave. egg Ave. no. CL of eggs (n) length (cm )eggs(n) width (cm; eggs(n) wt. (g) wt. (g) eggs(n) 17-18 9 (1) 2.78 9 (1) 1.79 9 (1) 53 5.89 9 (1) 18-19 19 (1) 19-20 9 (1) 3.67 9.5(2) 20-21 12.4(5) 3.24 11.3(3) 2.14 12 (1) 117 9.75 11.8(4) 21-22 14 (2) 3.38 17 (1) 147 8.65 13.3(6) 22-23 14.9(11) 3.31 14.8(6) 2.24 12.8(5) 134 10.5 14.0(16) 23-24 15.8(6) 3.45 16 (5) 2.32 13.6(5) 135(5) 9.90 14.8(10) 24-25 14.5(6) 3.48 14.2(4) 2.33 14.7(3) 147(3) 9.98 15.5(8) 25-26 16 (1) 3.88 16 (1) 2.47 16 (1) 26-27 22 (1) 3.50 22 (1) Mean 14.1 eggs/clutch taken from female G. oiiachitensis and G. pseiidogeographica. Examination of re- June and July 1972 suggests that females ordinarily lay two clutches of eggs per sea- son. This was determined by presence of two sets of corpora lutea, two sets of cor- pora iutea and shelled eggs, or one set of corpora lutea and a set of enlarged fol- licles. However, there was evidence to suggest the possibility of three clutches in eight individuals. The average clutch size of 50 G. pseii- dogeographica was 14.1 giving an annual reproductive potential of 28.2. Clutches of 65 G. oiiachitensis averaged 10.5 eggs for an annual reproductive potential of 21. These figures compare with those for G. piilchra, 29 (Shealy, 1976) and G. bar- boiiri, 17 (Cagle, 1952), both of which are slightly larger in carapace length. As in C. piilchra (Shealy, 1976) there was a direct correlation between clutch size and carapace length (Figs. 2 and 3); the long- er the female, the greater the number of eggs per clutch. Tables 2 and 3 show the relationship between mean number of eggs and mean egg length, mean egg width, mean egg weight and mean clutch weight at specific carapace lengths for both G. oiiachitensis and G. pseiidogeo- graphica. The mean clutch weight and mean egg length increase most noticeably with increase in both female carapace size and clutch size. Average egg weight and egg width are not so strongly correlated. Tables 4 and 5 show the relationship be- tween clutch size and egg weight, length and width. No direct trends or correla- tions are obvious. The method of estimating reproductive potential by counting corpora lutea, en- larged ovarian follicles and oviducal eggs has been used in many turtle studies (Cagle, 1952; Moll and Legler, 1971; Christiansen and Moll, 1973; Moll, 1973; Shealy, 1976). Legler (1960) describes the structures in detail. Although this method appears to be realistic, it may lead to overestimation or underestima- tion of annual reproductive potential. The presence of enlarged follicles toward the end of the laying season must be in- terpreted with caution. These follicles could represent clutches to be laid the following year, particularly if the indivi- dual had nested early the same year. 26 Tulane Studies in Zoology and Botany Vol. 22 ' Table 3. Relationship of both carapace length (CL) and number of eggs per clutch (n = number of clutches measured) and how they relate to size and weight of eggs in Graptemvs ouachitensis. Ave. no. Ave. egg Ave. no. Ave. egg Ave. no. Ave. clutch Ave. egg Ave. no. CL ofeggs(h) length (cm) eggs (n) width (cm) eggs (n) wt. (g) wt. (g) eggs (n) 17-18 11 (1) 3.02 8 (1) 74 9.25 9.5 (2) 18-19 9.3 (6) 3.28 9.2(4) 2.18 8.6(5) 82 9.54 9.3(7) 19-20 8.64(11) 3.46 9.2(7) 2.16 8.7(9) 81.8 9.4 8.5(14) 20-21 10.22 (9) 3.43 9.8(6) 2.16 9.7(14) 99.6 10.3 10.1(19) 21-22 13.4 (9) 3.36 12.6(8) 2.19 12 (10) 129.2 10.8 12.8(13) 22-23 12 (2) 3.44 12 (2) 2.24 13.3(4) 143.8 10.8 13.2(6) 23-24 11 (2) 3.47 11 (2) 2.12 14.5 (2) 153 10.5 12.3(3) 24-25 15 (1) 3.59 15 (1) 2.95 15 (1) Mean 10.5 eggs/clutch Eight females collected between 8-15 July contained old corpora lutea, ovidu- cal eggs and enlarged follicles. The latest date Graptemvs was found with oviducal eggs was 28 July, so these enlarged folli- cles possibly could have ovulated later that same season. One G. ouachitensis collected 8 July 1972, contained 12 new corpora lutea, 25 old corpora lutea and enlarged ovarian follicles. Because the average clutch size of G. ouachitensis is 10.5 and the record number of eggs per clutch is 17, the 25 old corpora lutea very probably represent two clutches. This fe- male laid three clutches with the potential of laying a fourth. Between 6 June 1972, and 13 August 1972, 641 mature G. ouachitensis and G. pseudogeographica were individually marked and released. No turtles were found to contain shelled eggs after they were known to have laid one clutch, but Table 4. Relationship of clutch size to egg weight, length and width regardless of the carapace size in G. ouachitensis (n = number of clutches measured). Clutch Ave. wt./ Ave. length/ Ave. width/ size eggs (g) n eggs (cm) n eggs (cm) n 6 9.17 3 3.34 2 2.11 2 7 10.00 2 — — — — 8 9.07 6 3.25 4 2.10 4 9 9.62 11 3.52 14 2.16 11 10 10.77 7 3.08 2 — — 11 12.18 6 3.33 8 2.15 8 13 13.43 3 3.46 1 2.22 1 14 10.64 3 3.40 5 2.21 5 15 9.73 1 3.28 1 2.95 1 16 11.63 2 3.70 1 2.26 1 479 47 39 34 eggs clutches clutches clutches No. 1 Wisconsin Map Turtles 27 since trapping did not start until 6 June, many of the turtles could already have laid their first clutch. Eggs were obtained from 16 G. pseu- dogeographica and G. oiiachitensis fe- males which were injected with oxytocin and released between 23 May and 2 June 1977. None of these was recovered during a five day trapping period, 9-14 June 1977 in an attempt to document the laying of more than one clutch in a season. How- ever, two G. oiiachitensis were observed to have double clutched in 1978. Twenty-four adult females collected on 15 October 1976, and maintained in the laboratory at 13^C were found to have atretic ovarian follicles upon dissection in January. The livers were distinctly marked with yellow suggesting eggs were being reabsorbed. Atresia has not been observed in freshly caught Graptemys during any season. Atresia of small or medium-sized follicles in Pseiidemys scripta in Panama was reported by Moll and Legler (1971) and by Webb (1961) in Oklahoma. Occurrence of atretic follicles is also known from several other species: Chrysemys picta (Powell, 1967), Terra- pene ornata (Legler, 1960) and Terrapene Carolina (Atland, 1951). The magnitude of atresia occurring in the 24 Graptemys maintained in the laboratory suggests that they are capable of complete atresia of all enlarged follicles if they are under environmental stress. NESTING Females emerge from hibernation and begin basking in early- to mid-April. The warmer the temperatures in April and May, the higher the metabolic rate of the turtles; thus the shell may be laid around the eggs more rapidly, allowing the clutch to be laid earlier. This was demonstrated by a female G. geographica collected at the Crosby Slough hibernaculum on 5 November and brought into the labora- tory and artificially hibernated at 3°-5°C until 10 January. She was gradually brought to 32^0 over a 24 hr period. Within 10 days shelled oviducal eggs were present. She was stimulated to oviposi- Table 5. Relationship of clutch size to egg weight, length and width regardless of the carapace size in G. pseudogeographica (n = number of clutches measured). Clutch Ave. wt./ Ave. length/ Ave. width/ size eggs (g) n eggs (cm) n eggs (cm) n 8 10.0 1 3.47 2 2.24 2 9 10 11 5.89 1 3.62 2 1.79 1 10.0 3 3.24 1 2.09 1 12 10.67 2 3.33 1 2.20 1 13 11.16 2 3.48 6 2.29 2 14 9.64 1 3.33 5 2.20 4 15 9.87 1 3.43 5 2.37 1 16 — — 3.74 2 2.47 1 17 8.65 1 3.24 1 2.16 1 18 10.0 2 3.44 3 2.28 3 19 9.36 2 3.23 2 2.20 2 183 16 30 19 eggs clutches clutches clutches 28 Tulane Studies in Zoology and Botany Vol. 22 tion by oxytocin injection on 25 January. After 2 months incubation at 25*^0 the eggs hatched. The onset of nesting ap- pears to be determined by spring tem- peratures. During the seven years of this study, the earliest first nesting observed was 18 May and the latest 10 June: the last known nesting occurred on 1 1 July. Females with shelled oviducal eggs have been found as late as 26 July (1972). Nesting was not observed in 1972 until 9 June, and 95*^ of the females caught af- ter 10 July did not contain oviducal eggs. This corresponds roughly with the length of laying period in 1977 when the earliest nest was recorded 18 May and laying was nearly finished by 14 June. Temperatures in April and May, 1977, were higher than in the same period in 1972, and may ac- count for the earlier onset of egg laying in 1977 than in 1972 (Table 6). Although the 1977 turtles conceivably could have laid a third clutch in late June or early July, none appeared to have done so since all eggs in the 128 nests examined on 19 Au- gust had hatched and only six had young remaining in the cavity. Map turtles have been found construct- ing nests from 0545-2030 CST under vari- ous weather conditions. The majority of nestings occurred between 0630 and lOCX). Later in the day, air and sand tem- peratures and sun intensity became too great for turtles to leave the water. Dur- ing overcast days nesting occurred throughout the day. Cool nocturnal air temperatures may also inhibit females Table 6 Temperature dal a ( C) from LaCrosse Weather Bureau (approximately 16 km north of study area) Month AP-^^I May June Year 1972 1977 1972 1977 1972 1977 Ave . Max . 11.7 19.0 24.6 26.1 25.2 26.0 Ave. Min. 0.6 6.2 10.3 13.2 11.9 n.o Ave . Temp . 6.1 12.6 17.5 19.7 19.2 20.0 from leaving the water to nest. Sixteen females were found on an overcast day at 1400 on Brownsville Island, South Beach in various stages from searching for dig- ging sites to completion of nests. A rain shower at 1430 stopped these activities, and all but one, which was covering her eggs with sand, began moving toward the water. She finished laying before return- ing to the water. Seven abandoned the nest cavities they had dug. Cloacal temperatures of turtles ob- served nesting ranged from 24.6°C to 28.2^C; air temperatures ranged from 21.10C to 32.0OC. During June, from 0430 to 1000 up to 75 females could be seen floating at the surface of the water 3 to 15 m from the shore of nesting beach- es. These turtles were apparently waiting for the right combination of environmen- tal cues before emerging. Once a female reached the beach she would wander from 5 to 150 m before attempting to dig a cavity. Nests were located in a variety of settings from the low shrubs surrounding the beach to the open sand area. Usually nests were dug adjacent to clumps of Ca- rf.vor other herbs. The nest cavities were dug entirely by alternate use of the hind limbs; no body pit such as that I have observed to be prepared by Chrysemys picta and Trionyx spiniferus, in this area, was dug. The nest cavity is flask-shaped and 10 to 16 cm deep; the neck broadens between four and eight cm beneath the surface. The eggs are positioned and packed into the hole with one of the hind feet. As an egg is being laid the turtle's neck is out- stretched fully and the head bobs slightly in the vertical plane. After the last egg is laid, the hole is filled by scraping and packing sand and nearby debris into the hole with the hindlimbs. Once the cavity is filled, the female continues to pack down the nest by alternately raising each hind limb 5 cm above the surface and then No. 1 Wisconsin Map Turtles 29 slapping the sand with it. The female leaves the nesting beach and returns to the water usually through the same break in the vegetation that was used in coming from the water. Fresh nests can be readily distinguished by the striking difference in sand particle size between the nest and the surrounding sand surface. The surrounding sand has a coarse mosaic of small pebbles created by wind erosion of the lighter sand while the nest has a rather uniform small-grained appearance. If found within an hour of nesting, the nest area is moist due to void- ing of bladder water and thus darker than the surrounding sand. Nests remain obvi- ous for a week or longer depending on rain and wind erosion. Settling of sand between recently laid eggs and after the eggs hatch forms a shallow crater making the nest even more easily distinguishable. Some of the nesting beaches are so crowded that nests may overlap. Areas where dredge spoil has been dumped in the last four years are not used for nesting. No plants grow on those sand piles and the water table is too low for the turtles to reach moist sand. Many at- tempts to excavate nests in dredge spoil banks were noted; digging was apparent- ly abandoned when the cavity failed to hold its form. Abandoned holes were also observed on other nesting areas. Other authorities (Legler, 1960; Shealy, 1976) refer to these as "test holes," implying that the turtle is testing the substrate be- fore laying. This may be the case in some instances, but it is also just as probable that the turtles were disturbed by weather conditions before egg laying commenced, and abandoned the nest. Turtles did not return to complete or lay in these aban- doned cavities. Nest temperatures were monitored continuously from 16 July to 17 August 1972 (Table 7). Thermistors were placed in the center of four nests. Two (a) were 10 cm beneath the surface and two (b) were 14 cm beneath the surface. Two ad- ditional thermistors were placed outside of nests: one 1 cm below the sand surface (c), the other 10 cm above the surface in a slatted wooden container shaded by a tree (Betula) (d). Daily temperature fluc- tuations ranged from 2.2 to 12.20C in the nests. The average daily fluctuations for two nests for 28 days was 6.7^0. The incubation period in nature ranged from 60-75 days (14 nests). Hatching success in nature was approximately 95% in 285 nests, (1972-77) that were excavated after hatching. Hatchlings usually remain in the nest until the yolk is completely ab- sorbed. Table 7. Nest, substrate and air mean temperatures (^C) from 16 July 1972 to 17 August 1972. Air in shade. Nest, 10 cm Nest, 14 cm Sand, 1 cm 10 cm above Air, LaCrosse below surface below surface below surface surface Weather Bureau Highest high 34.4 31.3 48.9 43.9 35.6 Lowest low 14.4 13.3 8.9 8.9 7.2 Average high 26.1 24.4 36.6 35.0 26.6 Average low 18.8 18.3 14.4 14.4 16.1 Daily mean 22.5 21.4 25.5 24.7 21.4 Daily fluctuations Largest 12.20C Lowest 2 .20C Average 6.7°C 30 Tulane Studies in Zoology and Botany Vol. 22 No positive evidence of hatchling over- wintering in the nest has been document- ed for any of the species at this site. Over 500 nests have been examined from mid- September through November without finding young which might yet overwinter before emergence. On 9 June 1972, one hatchling was found crawling across the center of nesting beach no. 5 at 0930 hours. The umbilical scar was dry and crusted and appeared as if the hatchling had not spent the winter underwater. This early sighting is the only evidence suggesting the possibility of occasional overwintering. Overwintering in the nest has been observed for other species Pseudemys scripta (Cagle, 1944), Chrys- emyspicta (Hartweg, 1946; Sexton, 1957; Ream, 1967; Vogt, 1978), and Graptemys spp (Newman, 1906). The head and limbs of the hatchlings of all three species are marked similar to those of the adults, but the carapace and plastron tend to be lighter and more bold- ly marked than in adult. Ontogenetic dif- ferentiation and pattern variation among species is discussed elsewhere (Vogt, 1978). Laboratory incubation One to two days prior to hatching in the laboratory, droplets of liquid condense on the surface of the egg. The egg by this time has expanded greatly in width and is reduced in length. Within 48 hours one or two longitudinal slits 5 to 20 mm in length appear. Presumably these are cut by the caruncle. These slits increase in length over the next few hours and the hatchling will force its head through the opening. In the laboratory hatchlings will often pull themselves out of the egg with their fore- limbs and bury themselves in the sub- strate. Usually, however, the hatchling will remain in the egg shell for 3-6 days until the yolk sac is nearly absorbed. The caruncle is shed or absorbed within three weeks of hatching. The temperature at which the eggs are incubated has a dramatic effect on head patterns, scute anomalies, and sex deter- mination. Each of 11 clutches of G. ou- achitensis and six clutches G. pseudoge- ographica were divided equally between incubators set at 25° and 35°C. Sixty-six eggs were incubated at 25°C, 54 hatched in 66-70 days, six died during develop- ment and six did not develop. Seventy eggs were incubated at 35*^C, 17 hatched in 49-54 days, 24 died during late stages of development and 29 failed to show any development. The low success at 35*^C can be attributed both to temperature and insufficient humidity. The eggs incu- bated at 35°C often became dented be- fore water was added to the substrate. Five percent of the 60 that developed at 25°C had scute anomalies while 29% of the 41 that incubated at 35°C had them. Another effect of incubation tempera- ture was the size of the yellow blotches on the head of both G. pseudogeographica and G. ouachitensis. Ewert (pers comm) noted that postorbital yellow "crescents" were formed in G. ouachitensis hatch- lings when eggs were incubated at low temperatures (25°C), but not at high temperatures (30°C). He did not notice this effect in G. pseudogeographica from Wisconsin. Measurement of the indivi- dual area of 13 of the yellow head blotch- es on hatchlings of both G. pseudogeo- graphica and G. ouachitensis showed sig- nificant differences between eggs of the same clutch incubated at 25°C and 35°C (Vogt, 1978). A greater percentage of larger yellow blotches were formed at the lower temperatures. Besides the phenotypic effects of incuba- tion temperature, the sex of the hatchling seems to be influenced by incubation temperature. All of those that hatched at No. 1 Wisconsin Map Turtles 31 25^C were male and all that hatched at 35°C were female. The probability that 54 of the 66 incubated at 25°C being male as a result of random sampling is very small (< .05 probability in Chi'- test). Bull and Vogt (1979) showed that sex is environmentally determined under both field and laboratory conditions. Yntema (1976) and Pieau (1971, 1972, 1973) sug- gested a similar phenomenon for Che- lydra serpentina and Emys orbicularis. The likelihood of the environmental determination of sex may help to explain the ratio of 5 females per male in the population. The collecting techniques used were sexually unbiased (Vogt, Co- peia, in press). Such skewed sex ratios have been reported for other populations of G. pseudogeographica (Timken, 1968). He found a 4:1 sex ratio in favor of females in the Missouri River in South Dakota. Hildebrand (1929) found a sex ratio of 4.4 females per male for captive raised Malaclemys terrapin. Most other turtle studies have found sex ratios which do not differ significantly from 1:1 (Gib- bons, 1970). My observations and collec- tions made of Graptemys ouachitensis and G.p. kohni in the White River, De- Vairs Bluff, Arkansas from 1975-1977 show the opposite trend. About four times as many males as females are seen and caught. Tinkle (pers comm) found the same to be true in Louisiana popula- tions of these species. The shorter testes can be distinguished at hatchling from the ovaries by the abs- ence of oviducts. Ovaries also have small white spots on the surface. The sex of a turtle apparently is determined prior to hatching, since no sex reversals have been noted from 30 hatchlings main- tained in the laboratory for nine months. Hatchlings that were maintained in the laboratory for 5.5 years showed no ten- dency toward sex reversal once secon- dary sexual characters became apparent. Growth in laboratory reared hatchlings So few yearling and immature turtles were discovered in the study area that hatchlings had to be reared under artifi- cial conditions to answer several ques- tions concerning growth and demograph- ic phenomena, such as: What is the sex ratio at hatching? Is the differential sex ratio in the adult population an index of the sex ratio at hatching or is differential mortality occurring? Do male turtles grow more slowly than females and thus are they more susceptible to predation for a longer period of time, or do male and female turtles grow at equal rates initially, following which males slow in growth upon reaching a specific size (sex- ual maturity)? When do the external sex- ually dimorphic characters appear? When is sexual maturity reached? A total of 197 hatchlings consisting of 65 G. ouachitensis from 17 clutches and 132 G. pseudogeographica from 19 clutches were reared in the laboratory. At hatching both species were of similar size (Table 8). Growth of 86 turtles was measured through 10 March 1975, but 47 of these died in the fall of 1975 due to a virulent infection of the bacteria Citrobacter freundii. The 39 that survived after treat- ment with Chloromycetin were main- tained through 25 October 1978. During the first three years males and females increased in mass and shell prop- ortions at nearly the same rate (Figs. 4-9). Male growth began tapering off between March 1975 and June 1976, at 3-4 years of age. Females, however, continued grow- ing up to the present. Table 9 summarizes the change in the mean monthly increment of growth for males and females in 1975 and 1978 and the effect these differences had on total mass and shell measurements. Monthly increase in carapace length of males and 32 Tulane Studies in Zoology and Botany Vol. 22 Table 8. Measurements of hatchling G. ouachitensis and G. pseudogeographica in- cubated in the laboratory in 1972. (mean above; range below) Species # of clutches # of hatchlings Ave. wt. (g) Carapace length (cm) Carapace width (cm) Plastron length (cm) Plastron width (cm) G. ouachitensis C pseudogeographica 16 10 116 99 4.4 g 4.99 g (1.5-6.2) (1.8-6.6) 3.07 3.04 (2.71-3.44) (2.50-3.30) 3.01 3.04 (2.65-3.40) (2.66-3.37) 2.85 2.84 (2.53-3.18) (2.38-3.16) 1.97 1.96 (1.75-2.65) (1.55-2.12) females was similar through March 1975, 0.50 mm and 0.43 mm respectively. But from March 1977, to April 1978, the aver- age growth per month differed dramati- cally, 0.56 mm for males and 1.6 mm for females. The average monthly increase in carapace length from November 1972, through March 1975, (27.5 months) was similar to males and females (1.27 mm and 1.38 mm). However, females grew at over twice the rate of males for the next 37 months (March 1975, through April 1978). Carapace length increased an av- erage of 1.06 mm per month in females, but only 0.52 mm per month in males. Reduction in growth rate of males coin- cided with the onset of sexual maturity. When the males were four years of age they were considered sexually mature on the basis of extensive development of the external secondary sexual characters (elongated 2nd, 3rd. and 4th foreclaws, elongation and thickening of the tail, and courtship and copulation behavior). By April 1978, none of the 1972 hatch- ling females had yet reached sexual ma- turity. This was evident by their being smaller than the smallest mature female found in the wild and possession of no palpable shelled eggs. At least two of the Table 9. Growth (cm) rates of 1972 hatchlings. Graptemys pseudogeographica and G. ouachitensis are combined - growth rates were the same. W March 1975 15 April 1978 Monthly Monthly Average Growth Average Growth 53 c? 33 9 53 05* 339 20 d" 19 9 20cf 199 CL 6.30 6.87 .50 .43 8.46 11.73 .56 1.6 CW 5.71 6.01 7.04 9.52 CH 2.68 2.79 3.37 4.87 PL 5.61 5.80 7.34 10.50 PW 4.10 4.20 5.12 7.07 Wt. 35.5 44.5 .96 2.52 85.0 268.4 1.07 7.69 No. 1 Wisconsin Map Turtles 33 /? 250 / 200 / 150 / 100 : y_^y-—"^ 50 ^^^/^--^""^ 1 >— ^°*^l II III NOV DEC APR MAY JUN MAY APR 72 73 74 75 76 77 78 Figure 4. Weight changes in 197 captive raised hatchlings. (65 G. ouachiiensis: 132 G. pseiido- geographica). G. pseudogeographica females will reach reproductive size by spring 1980 if pre- sent growth rate continues. Because of the continuous warm temperatures and high intensity light regime under which the turtles were raised and continuous NOV DEC APR MAY JUN MAY APR 72 73 74 75 76 77 78 Figure 6. Carapace width changes in captive raised hatchlings. feeding, these laboratory populations cannot be compared directly to natural ones. However, females did become less active and ate less from January through March each winter when ambient exter- nal temperatures reduced the interior vivarium temperatures approximately 70c. Growth rings The use of growth rings to estimate the age of turtles has been reported by nu- Figure 5. Carapace length changes in captive raised hatchlings. Figure 7. Carapace height changes in captive raised hatchlings. 34 Tulane Studies in Zoology and Botany Vol. 22 ' Figure 8. Plastron length changes in captive raised hatchlings. merous authors {Graptemys pulchra, Shealy, 1976; Terrapene ornata, Legler, 1960; Pseudemys scripta, Cagle, 1946; 1950; Moll and Legler, 1971; Chrysemys picta. Sexton, 1959; Gibbons, 1967; Ernst, 1971; G. ouachitensis, Webb, 1961; Pseudemys rubriventris, Graham, 1971; Gophenis agassizi, Patterson and Brattstrom, 1972). Counting of growth rings gives a rough estimate of the age of the turtle, but can be used reliably only if the areola (hatch- ling scute) is present. In the Stoddard population of Graptemys (all 3 species) the maximum number of rings observed is six in males and up to twelve in females (Table 10). After that time earlier rings become worn off, and growth is so limited that new rings are difficult to distinguish. As the turtles continue to shed epidermal scutes the rings become less and less visi- ble, until the shell is relatively smooth. Thus reliable estimate of the ages of most males over 6 and females over 12 years was impossible. A Graptemys pseudoge- ographica survived 32 years captivity (Bowler, 1977). Cagle (1950) suggested that Pseudemys scripta elegans were re- productively active for 40 to 50 years, and that natural longevity extended from 50 to 75 years. Moll and Legler (1971), how- ever, were more conservative, estimating 30 years longevity for tropical Pseudemys scripta under natural conditions. Gib- bons (1968) estimated some Michigan Chrysemys picta to be nearly 40 years old. Judging from the size and age structure of the Stoddard population, and the degree to which shells of some individuals are worn, 30-50 years seems a reasonable es- timate for the present age of many indivi- duals. Natural Growth The smallest sexually mature male G. ouachitensis caught in the wild was 7.48 cm in carapace length. Wild caught G. geographica, G. ouachitensis and G. pseudogeographica males were not re- productively mature in their 3rd year, but did mature in four to six years. gov 72 DEC APR 7 3 74 MAY 75 JUN 76 MAY 77 APR 78 Figure 9. Plastron width changes in captive raised hatchlings. No. 1 Wisconsin Map Turtles 35 The youngest mature female G. ou- achitensis and G. pseudogeographica, as evidenced by the presence of shelled eggs, were in their eighth year. Female G. geographica that were in their tenth and twelfth years had not yet matured (based on a lack of enlarged ovarian fol- licles). Figures 10 and 11 and Table 10 show the relationships between the number of growth rings and the weight and carapace length respectively for G. ouachitensis fe- males from the wild. The growth rate pat- tern of wild females is similar to that shown for captive raised hatchlings (Figs. 4 and 5). The curve (Fig. 10) shows a trend in which carapace length levels off and growth slows. Like the males, fe- males appear to level off in carapace growth once reproductive maturity is reached. Recapture of nine G. pseudogeo- graphica and six G. ouachitensis adult fe- males between one and five years after first marking showed very little increase in carapace length, 1.1 mm per year (Table 11). Two individuals decreased slightly in length, which could be due to carapace wear or original measurement error or shrinkage. Ream (1967) also re- ported loss in carapace length in Chrys- emys picta. Although increase of the carapace length slows after maturity, 13 individuals exhibited an increase in body weight. The average change in weight for the 15 females (both species) was +49.2 g/yr. Laboratory raised females gained nearly twice as much in body weight in their fifth year of growth, 96.06 g/yr. Changes in carapace cross section to ac- commodate more eggs continue after ma- Table 10. Number of growth rings and size of wild caught G. ouachitensis'^ s. Age Wt. (g) CL (cm) CW (cm) CH (cm) PL (cm) PW (cm) Maturity 0 4.4 3.1 3.0 1.4 2.9 2.0 hatchling 2 53 7.6 6.5 3.2 6.7 4.5 juv. 3 64 7.8 7.0 3.5 7.3 5.4 juv. 4 170 10.9 9.4 4.6 10.2 6.8 juv. 5 261 12.5 10.2 5.2 11.8 7.5 juv. 6 515 16.6 12.8 6.8 15.8 9.5 juv. 7 441 15.7 12.2 6.2 14.4 9.3 juv. 8 572 16.6 12.6 6.7 15.7 9.4 8 605 16.5 13.4 6.7 16.2 10.7 8 767 18.3 14.2 7.4 17.2 11.0 8 564 16.3 12.8 6.7 15.2 9.3 8 560 17.2 13.2 6.5 15.4 9.7 8 557 16.7 13.1 6.2 15.6 9.3 9 983 20.3 16.1 8.1 19.3 12.2 9 748 17.8 13.6 7.6 16.6 10.2 eggs 7 1000 16.3 eggs 9 682 17.4 immature 9 555 16.0 immature 10 1031 20.0 15.3 8.0 18.2 11.3 10 815 19.1 14.1 7.4 17.7 11.1 11 714 17.4 13.8 7.2 16.1 12.4 eggs 11 1048 21.1 16.5 8.5 19.3 12.0 eggs 12 1032 20.0 15.7 8.4 18.8 11.7 36 Tulane Studies in Zoology and Botany Vol. 22 22 20- 18- 16 14- G ouochitensis ? < Q. ^ 12 < 10- 8- _ • • _ • • • • • • • ! • _ • • ~ • • 1 1 1 1 1 1 1 1 1 1 3456789 10 II NUMBER OF GROWTH RINGS 12 13 Figure 10. Relationship between carapace length and number of growth rings in wild caught fe- male G. ouochitensis. turity and probably are responsible for the increase in weight. Males show little change in carapace length or shape fol- lowing maturity. POPULATION COMPOSITION Frequencies of size classes of carapace length and weight are shown in Figures 12-15. Definite peaks in both weight and carapace length depict the larger average size of G. pseudogeographica females; 82% of the G. pseudogeographica fe- males range between 21-25 cm while 85% of the G. oiiachitensis females range be- tween 19-23 cm. G. oiiachitensis also weigh less than G. pseudogeographica females: 85% of the G. ouochitensis were between 800-1400 g while 79% of the G. pseudogeographica were in the 1100- 1800 g categories. The peak carapace length reached in females probably rep- resents the size at which growth rate de- creases at the onset of reproductive ma- turity. The broader distribution for weight may be due to bias introduced by weighing turtles with full or empty blad- ders, or with or without mature eggs. Males of both species are extremely uniform in weight and body size. This reflects the early age of maturity and the slowing of growth after maturity, as was shown in the laboratory population. POPULATION DYNAMICS The size of the study area and the mo- bility of the turtles did not allow a reason- able estimate of population density through traditional mark and recapture techniques. Individual turtles were found to move over 4 km up and down stream within the course of a year, so the use of 6 0 uochitensis -f- 1000 3 45 6 78 9 10 II NUMBER OF GROWTH RINGS 12 13 Figure II. Relationship between weight and number of growth rings in wild caught male G. ouochitensis. No. 1 Wisconsin Map Turtles 37 traditional mark and recapture index would tend grossly to overestimate the number of turtles actually present. Even in the fall when they are at the overwin- tering site turtles are moving to different locations (Table 12). This was evidenced by the capture and release of 131 Grapt- emys on 13 October 1977 at the "hiber- naculum" in Crosby Slough. On 11 No- vember 1977, 113 G raptemys yfjQXQ cdiXighi at the same site, but only two of these were turtles that were marked on 13 Oc- tober. From 3 June through 13 August 1972, stationary fyke nets were set, and usually checked daily, at 15 localities at Stoddard (Fig. 1). During that period, 326 G. pseu- dogeographica (299 females and 27 males) and 168 G. ouachitensis (167 fe- males and one male) were captured. The highly skewed sex ratio apparent in these totals was biased by extensive trapping adjacent to nesting beaches where females were usually abundant. Traps adjacent to the beaches caught 257 females and only two males. Traps set away from beaches and sloughs, in shal- low weedy areas caught 196 females (156 G. pseudogeographica, 40 G. ouachiten- sis) and 26 males (25 G. pseudogeograph- ica, one G. ouachitensis). This was still an overestimate of females since male G. ouachitensis were found primarily along the sloughs in moving water. Female G. ouachitensis were found primarily in the backwater areas adjacent to islands and Table 1 1 . Growth of marked turtles in the wild. # CL (cm) A CL (cm) A Wt. (g) Time after marking Female G. pseudogeographica 256 22 .9 (.225) 158 (39.5) 4yr. 342 23.1 .2 (.047) 613(144.2) 4yr.,3mo. 441 23.8 .1(.044) 152 (67.5) 2yr., 3 mo. 57 23.1 .4 (.092) 215 (49.88) 4 yr., 4 mo. 59 24.8 2.0 (.4) 412(82.4) 5yr. 204 21.0 0 102(19.4) 5 yr., 4 mo. 331 22.9 .2 (-.041) 425 (86.7) 4yr., 11 mo. 418 23.1 .7 (.143) 180 (36.7) 4yr., 11 mo. 594 22.8 1.0 (.02) 58(11.8) 4yr., 11 mo. Male G. pseudogeographica 595 -11 4yr., 11 mo. 598 15.1 .6 (.12) +23 5yr. 602 13.4 .1 +6 4yr., 11 mo. Female G. ouachitensis 214 21.8 .8 (.2) 297 (74.25) 4 yr. 685 20.9 • K.i) 53 (53) lyr. 714 19.2 0 43 (-43) lyr. 496 22.1 .3 (.071) -15 (-3.53) 4 yr., 3 mo. 47 20.8 1.8 (.45) 250 (62.5) 4yr. 377 20.6 (-.l)(-.044) 128 (56.8) 2 yr., 3 mo. CL = carapace length at last capture ACL = change in carapace length since first capture with average yeariy change in parenthesis;AWt. = change in weight since last capture with average yearly eain in oarenthesis 38 Tulane Studies in Zoology and Botany Vol. 22 % 100 n - 488 G pseudogeographica * i40r 120- 100 0 pieudogeogrophico i 600 1000 1500 WEIGHT (q) 2000 2500 0-15 16 18 20 22 24 26 28 CARAPACE LENGTH (cm) Figure 12. Weight and carapace length classes of G. pseudogeographica females captured. sloughs and were less abundant in shal- lows away from islands or in moving wa- ter. Gill nets set by commercial fishermen adjacent to the nesting beaches of the main island caught nearly equal numbers of female G. ouachitensis (44) and G. pseudogeographica (45), but more fe- male G. pseudogeographica (142) than G. ouachitensis (1 15) were caught in fyke nets adjacent to the nesting beaches. Fe- male and male Graptemys ouachitensis outnumbered G. pseudogeographica when trammel nets were set in the sloughs near overwintering sites. In the fall of 1976 and again in 1977, 239 G. ouachitensis (191 females and 48 males) and 67 G. pseudogeographica (45 females and 22 males) were captured. Since males of both species are found primarily in the G ouachitensis ? 0-500 i60r n = 521 1000 1500 WEIGHT (g) 2000 0-15 16 18 20 22 24 CARAPACE LENGTH (cm) Figure 13. Weight and carapace length classes of G'. ouaclutensis females captured. No. 1 Wisconsin Map Turtles 39 Table 12. Turtles caught at Crosby Wing Dam overwintering site. Date Oct. 1. 1976 Oct. 13. 1977 Nov. 11. 1977 G. geographica G. ouachitensis G. psendogeographica Cf 9 Juv. cf 9 Juv. cf 9 Juv. Total 0 4 0 7 49 0 6 3 0 0 30 67 1 14 9 5 1 11 70 4 1 48 186 IT 19 13 11 43 0 1 1 85 131 113 329 sloughs and female G. pseudogeograph- ica occur primarily in the backwaters, male G. pseiidogeographica are over-rep- resented in this sample. The ratio of male to female G. ouachitensis, 1:4 is fairly realistic but may be an overestimate of the males. In 1972 a total of 802 G. ouachitensis and G. pseudogeographica was captured. The sex ratio of 12.6 females per male was biased due to trapping near nesting beaches and use of gill nets. An addition- al 315 (236 females and 79 males) were captured at intermittent trapping inter- vals in 1975 and 1977. Graptemys ouachi- tensis and G. pseudogeographica make up nearly equal numbers of the 1,117 tur- tles sampled, 573 and 544 respectively. Graptemys geographica were not marked in 1972, but were less abundant, making up less than 10% of the total number of Graptemys captured. In 1976 and 1977 this species made up only 1% of the total number caught at the hibernaculum. Trapping results from 3 June through 13 August 1972 ranged from a daily catch of 106 on bright, warm, sunny days dur- ing the height of the nesting season to none on cold, rainy days. Fyke nets were used a total of 45 days, capturing an aver- age of 1 1 new individuals per day. Only 48 turtles were recaptured one or more times during the course of the study. Any turtle recaptured the day after G pseudogeographica Q, 0 200 400 WEIGHT (g) 10 12 14 16 CARAPACE LENGTH (cm) 50 < "Oh Z) Q o 20 C ouachitensis (5 10- n 67 I 1 /// ^^^ M-1 . P^ i 1 1 0 200 400 WEIGHT (g) 8 10 12 14 CARAPACE LENGTH (cm) Figure 14. Weight and carapace length classes of G. Figure 15. Weight and carapace length classes of G. pseudogeographica males. ouachitensis males. 40 Tulane Studies in Zoology and Botany Vol. 22 release was not recorded since release was made near the trap site increasing the probability of it being caught in the same trap as it attempted to escape. MOVEMENTS During 1972 marked turtles moved dis- tances as far as 4 km. Recapture intervals for females adjacent to nesting beaches ranged from three to 11 days. Since all of these had shelled oviducal eggs upon first capture I assumed that they stayed near the beaches in preparation for laying. Turtles which were caught near the beaches without shelled eggs were never recaptured in the same area in 1972. Twelve were recaptured between 0.4 km and and 1.6 km north of the island from three to 41 days later. Once a female had laid her eggs she vacated the area adja- cent to the island and moved into a regu- lar feeding area. Sixteen turtles were re- captured in the same backwater feeding areas 4 to 35 days apart suggesting a regu- lar home range. One female G. pseudo- geographica first caught after nesting was recaptured three days later 1.6 km north of the island. Three gravid females caught near the island in 1972 were recaptured near the same beaches in May, 1977. This suggests that an individual turtle may return to the same area to nest year after year. Two females marked near the nesting beaches in June, 1972, were found in October of the same year 4 km north in Crosby Slough in a communal hibernaculum. Nine more recaptures from 1972 were made there in 1976 and 1977. Two turtles hibernating in Crosby Slough in 1976 were recaptured there in fall 1977. A female G. pseudogeographica found on the nesting island 14 June 1976, was released at the boat landing at Stoddard, 15 June 1976. She was recaptured at the Crosby Slough hibernaculum 13 October 1977. This is approximately 8 km from her point of release. Shealy (1976) showed that female G. pulchra were cap- able of homing up to 15 km both up and downstream from points of release to points of original capture, but displaced males tended to be recaptured near the release point. He suggested that this could be because only males have a de- fined home range. A male G. pseudoge- ographica caught at Crosby Slough wing dam 14 October 1972, was recaptured at the same place 13 October 1977. It had been released at the Stoddard landing in May, 1973, so, like females, males pos- sess homing ability; however, if males regularly travel long distances is not known. The long distances regularly moved by the females from overwinter- ing areas, to nesting grounds, to feeding areas and back to overwintering areas suggests that females have a large irregu- lar activity area defined by the pool itself. FOOD AND FEEDING Detailed analysis of food partitioning and feeding behavior is reported in Vogt (1978). The stomach contents taken dur- ing June, July, and August from 21 fe- male G. geographica, 54 female G. ou- dchitensis and 35 female G. pseudogeo- graphica were quantitatively analyzed by volume. Female Graptemys geographica were shown to be mollusk specialists, mollusks made up 66% by volume of all their food. Insect larvae and fish carrion comprised most of the remainder. Mol- lusks were also important in the diet of female G. pseudogeographica (19% by volume). In G. ouachitensis , however, mollusks made up only 2.8% of the vol- ume. Vegetation comprised only 3.9% of the volume of material consumed by G. geo- graphica, whereas G. ouachitensis and G. pseudogeographica consumed 31.5% and 42.4% respectively. Algae, Potamoge- ton, Lemna and Vallisneria were the No. 1 Wisconsin Map Turtles 41 plants most frequently eaten. Insects (caddisfly cases, mayfly larvae, damselfly larvae) comprised 519^ by vol- ume of foods eaten by G. oiiachitensis females; while only 21.9*"^ and 15.3'^f in C pseiulogeographica and G. geograph- ica respectively. No differences in food habits among the males of the three species could be determined. Mayfly larvae, damselfly larvae, caddisfly cases, beetles, flies, other insect larvae, moUusks and fish car- rion were eaten by males of all three spe- cies but only trace amounts of vegetation were found to have been eaten. ACTIVITY Seasonal Map turtles become active in western Wisconsin in April when they begin dis- persing from their overwintering sites while the water temperature is still A-l^C (Fig. 16). Mating presumably takes place while the turtles are still concentrated around the hibernaculum. but due to wa- ter opacity, courtship has never been ob- served in the wild. I have observed spe- cies such as Emydoidea hlandingi, Chrys- Seosonal Activity Patterns Fiiiurc 16. Generalized seasonal activity patterns i)t' Ci. Oiiachitensis and C pseitdogeograpfiica. 1. Emergence from overwintering sites. 2. Disper- sal from o\ervvintering sites and basking. 3. Egg laying. 4. Feeding. 5. Hatchling emergence. 6. Backing and movements to overwintering sites. 7. Agereeatine at overwintering sites. emvs picta and Chelydra serpentina mat- ing and courting at this time both north and south of this site in Wisconsin, thus water temperatures are not too cold to allow turtles to breed at this time. During May females move out of the channels and toward the nesting islands. They concentrate to bask by the dozens on the few available emergent logs near the nesting beaches. Male G. ouachiten- sis remain along the channels and sloughs, but during the summer many male G. geographica and most male G. pseiidogeographica move into the quiet backwaters to forage. This is evidenced by the summer fyke net catches in 1972, in which 27 male G. pseiidogeographica and two male G. oiiachitensis were cap- tured in the shallow, weedy area, 0.4 - 2.4 km north of the island. Many male G. geographica were also caught in this area, but were not marked. Earliest documented feeding by fe- males occurred on 26 May. There are several possibilities why feeding does not commence until this time: (1) there is not enough room in the body cavity for the stomach or the intestine to expand with food until the first clutch of eggs is laid; (2) the aquatic vegetation and insect lar- vae on which they feed have not grown to harvestable quantities; or (3) tempera- tures are not high enough to allow effl- cient digestion. All of these ideas are plausible for fe- males of both G. oiiachitensis and G. pseiidogeographica, but females of G. geographica feed nearly exclusively on mollusks which are abundant throughout the year. They do not consume as large a volume of food as do the other two spe- cies, so do not have as great a problem with limited gut space. The earliest G. geographica with food in its stomach was located 26 May, only 4 days before the first G. pseiidogeographica and G. oiia- chitensis. Why feeding is postponed until 42 Tulane Studies in Zoology and Botany Vol. 22 such a late date is unclear. In late May or early June (23 May-10 June) the turtles lay their first clutch of eggs and then move away from the nest- ing islands to feed for two to three weeks while shell is being deposited around the second clutch of enlarged ovarian folli- cles. Graptemys geographica and G. pseu- clogeographica move far into the back- waters where there are patches of Pota- mogeton and Vallisneria and many stumps available for basking. Though some females of G. oiiachitensis also move into the backwater areas, many stay near the nesting islands to feed and bask or move to the vegetation patches immediately adjacent to the river chan- nel. After laying their second clutch of eggs the females return to the feeding grounds. Most female G. oiiachitensis then move away from the nesting islands to the vegetation patches adjacent to the sloughs or channels. Feeding continues until the first week of September. The turtles then spend the next two months basking, during which time their guts are evacuated presumably so that they can enter hibernation with an empty gut cavity. Turtles which were not allowed to clear the gut cavity before they were artificially hibernated in the labor- atory died in 1-2 months, whereas those that were allowed to, lived until Spring. By October map turtles begin moving to the wing dams along the sloughs and channels. Here they are able to lodge themselves within the rock piles, so not to be carried away by the current and are supplied with a constant flow of well-ox- ygenated water. They are not totally dor- mant throughout the winter as commer- cial fishermen occasionally catch them in their nets while gill netting under the ice. The seasonal activity patterns of G. ou- achitensis and G. pseudogeographica are generalized graphically in Figure 16. Basking activity The functions of basking behavior in turtles have been discussed at length by other investigators (Cagle, 1950; Boyer, 1965; Moll and Legler, 1971 ; Auth, 1975; Shealy, 1976). The primary function of basking is thermoregulatory in nature. On 14 June 1977, between 1545 and 1550 hours, six female G. oiiachitensis and one male G. oiiachitensis were chased from a basking log into a trammel net. The air temperature (sky partially overcast) was 21.5^C and the water, 22.5°C. Cloacal temperatures of the six females ranged from 23.8°C to 27°C (mean 25.8°C); the male was the same temperature as the water - 22.5°C. No other data were obtained on internal temperatures of basking turtles. Since they bask for hours even at ambient tem- peratures above 350C, on sunny days their body temperatures must reach con- siderably higher. Moll and Legler (1971) reported voluntary core temperatures in basking Pseiidemys scripta as high as 35.7°C. As temperatures approach the critical thermal maximum, basking tur- tles begin gaping (Moll and Legler, 1971). Male Graptemys were observed gaping more frequently than females. Basking activity was observed between 0900 and 1800. Throughout the day a tur- tle would alternate between basking, feeding or other activity in the water. Basking sites were at a premium in the study area. Turtles used partially emerged logs, stumps, rock piles, musk- rat houses, and sand bars as basking sites. Logs or stumps that were not adjacent to a shoreline were used more frequently than any other type of basking site. Many, particularly males, would often climb onto the highest limbs of a basking log, often 2 m above the water. As turtles climbed onto basking logs they would usually become oriented in an anterior to posterior or posterior to posterior direc- No. 1 Wisconsin Map Turtles 43 tion with respect to nearest neighbor. Not only would this reduce the possibility of aggressive interactions by avoiding head to head contact, but also allowed for clos- er packing on the log by allowing a turtle to crawl partially upon the carapace of another turtle. I did not observe any tur- tles basking with the plastrons resting over the anterior end of the carapace of another turtle. Up to 47 adult Graptemys were observed basking simultaneously on a 6 m log with no aggressive interactions. Chrysemys picta belli, Trionyx muticus and T. spiniferus were also observed us- ing the same basking logs as the Grap- temys. Male T. spiniferus were observed biting male Graptemys that moved into the proximity of the head region on two occasions. Basking turtles were more prone to dis- turbance by passing boats within the first 5 min of emergence. The diving of one turtle into the water often triggered the entire log of turtles to slip in. After look- ing in several directions the turtles would all resume basking within 5 to 10 min. Individuals were noted to remain on a basking log for over 100 min before being disturbed. They occasionally lost their balance and fell into the water, precipi- tating sudden evacuation of the entire log. Throughout the time Graptemys are basking they are extremely alert to and wary of the slightest disturbance. Ap- proach within 30 meters without disturb- ing them is difficult unless the observer is snorkeling in the water. Communal bask- ing may be advantageous for the sighting of potential predators and also attracting grackles. Once a turtle became stationary on a basking log it would stretch out the hind- legs to their fullest extent and spread the webbing between the toes. The neck and forelimbs would also be extended. Pre- sumably this position functions to dry out the skin and increase the surface area exposed to solar radiation. But it also has another purpose; to expose the areas un- der the limbs where leeches attach. Com- mon grackles (Quiscalus quiscula) were observed on 10 June and 14 June remov- ing leeches (Placobdella parasitica) from the leg and neck cavities of basking Grapt- emys (Vogt, 1979). The grackles moved along the basking log inspecting each tur- tle, often walking around inspecting all leg cavities of an individual turtle. The turtles seemed to be oblivious of the grackle, even when the turtle was rocked back and forth on the log by the bird pulling at a particularly tenacious leech. Both male and female turtles allowed grackles to hunt for leeches on their bod- ies. The only two instances of grackle grooming were on basking logs contain- ing over 20 adult turtles. At what size turtles no longer view the grackle as a potential predator and allow cleaning would be interesting to learn. Predation No predation upon basking turtles was noted. Nests and eggs were destroyed by red fox (Vulpes fulva), raccoon (Procyon lotor), and otter {Liitra canadensis). Most {90^c ) of the nests on beaches 2 and 3 were destroyed within 24 hours after laying. If the nest was not raided within 48 hours it was usually not bothered. During 1977 raccoon predation was responsible for the destruction of over 90% of the nests on both ends of Browns- ville Island. No nest predation was ob- served on the main nesting island in 1976 or 1977. Otters excavated nests and con- sumed eggs during June, July, and Au- gust. Adult map turtles appear to have few potential predators in the study area. In August, when the hatchlings move to the water, flocks of ring-billed gulls (Larus delawarensis), crows {Corvus brachyrhynchos), grackles, and red- winged blackbirds (Agelaius phoenice- 44 Tulane Studies in Zoology and Botany Vol. 22 us), have been observed covering the beaches. On one occasion crows were found excavating a nest where hatchlings were emerging. Redwinged blackbirds and grackles were observed eviscerating hatchlings, leaving only the shells on the beach. Great blue herons {Ardea herodi- as) were also seen walking on the nesting beaches from 0600-0700, when hatchlings were leaving the nest, but no predation was observed. Many turtles are found with the carapace gashed, presumably by outboard motors. Individuals missing one forelimb, two forelimbs, or one hind- limb seemed to be able to function ade- quately, and were found laying eggs. An adult female G. ouachitensis collected in 1972 had recently suffered severe injury to the anterior part of the carapace and had lost half of the mandible. She was recaptured in 1976 still able to function with only half the mandible, and her shell was not completely healed after four years. SUMMARY Detailed analysis of the systematics of the false map turtle complex indicates that Graptemys ouachitensis and Grapt- em\s pseudogeographica are separate species (Vogt, 1978). Natural history data were gathered from populations on the Mississippi River, Vernon County, Wisconsin from 1972 through 1978. Tur- tles were collected in a 230 hectare area by using fyke nets with leads, gill nets, trammel nets and hand captures. Clutch- es of eggs were collected from females and incubated in the laboratory under various temperature regimes. Hatchlings were maintained in the laboratory for six years to study differential growth rates of males and females, and to determine the time at reproductive maturity and the sex ratios at hatching. Examination of reproductive tracts of 50 females suggests that they annually lay two clutches of eggs. Eight females per- haps laid three clutches. The average clutch size of 40 G. pseudogeographica was 14.1 and for 65 G. ouachitensis was 10.2. There is a direct correlation be- tween clutch size and female carapace length. The mean clutch weight and mean egg length increase with an increase in female carapace length and also the clutch size. Females begin developing enlarged ovarian follicles in mid-July and enter hi- bernation with the body cavity packed with enlarged follicles in October (Fig. 16 shows the occurrence of major seasonal activity patterns). Mating probably oc- curs either in October or April while the turtles are congregated around hibernac- ula. The courtship displays of male G. ou- achitensis and G. pseudogeographica in- volve drumming of the foreclaws against the ocular region of the female. There was a mean of 10.3 (n = 26) contacts per "titillation" bout for G. pseudogeograph- ica and only 5.2 (n = 24) for G. ouachi- tensis. Graptemys pseudogeographica al- so differs by bobbing its head in a vertical plane while G. ouachitensis holds its head stationary while vibrating the foreclaws. Fresh nests were found from 18 May to 1 1 July, but females with shelled oviducal eggs were found as late as 26 July. How- ever, 95% of females captured after 10 July lacked oviducal eggs. Nest temperatures, monitored contin- uouslv from 16 July to 17 August 1972, fluctuated from 2.20C to 12.20C daily. The mean daily fluctuation for two nests for 28 days was 6.70C. Natural incuba- tion periods ranged from 60-75 days. Hatching success is approximately 95% in both laboratory and nature. Incubation temperature in the labora- tory was shown to affect hatching success, number of scute anomalies, size of yellow blotches on the head, and sex determina- No. 1 Wisconsin Map Turtles 45 tion in both G. ouachitensis and G. pseu- dogeographica. More scute anomalies oc- curred at 350c than 250C. The yellow blotches on the head were larger on sibl- ings incubated at 25^0 than on those at 350c. All eggs that hatched at 25^0 (54 of 66) were male. All eggs that hatched at 35^C (n = 17) were female. These results strongly suggest that, at least in the lab- oratory, sex can be determined by early developmental temperatures. A sex ratio of 5 females per male was calculated from trapping results. Sex determination by developmental temperature may help to explain the skewed sex ratio in both G. ouachitensis and G. pseudogeographica. Sixty-five G. ouachitensis and 132 G. pseudogeographica hatchlings from 1972 were raised in the laboratory. During the first three years male and females (n = 86) increased at nearly the same rate in mass and shell proportions. Male growth slowed between March 1975 and June 1976 while females continued to grow at the initial rate. Males attained sexual ma- turity in their fourth year, as evidenced by secondary sexual characteristics, and the slowing of their growth rate is appa- rently correlated with this. Wild male Graptemys of all three species were ma- ture when four to six years old. No females had matured sexually by April 1978. The youngest wild-caught mature female G. ouachitensis and G. pseudogeographica were in their eighth year. Growth rings were used reliably to estimate age only during the first six years in males and up to 12 years in females. Mature male and female G. pseudogeo- graphica and G. ouachitensis recaptured after one to five years showed little in- crease in carapace length (less than 1.1 mm per year). In 1972 a total of 802 G. ouachitensis and G. pseudogeographica were cap- tured. Both male and female G. pseudo- geographica are able to return to their site of original capture after being displaced downstream as far as 8 km. The long dis- tances regularly moved by females from hibernating sites to nesting grounds, to feeding areas and back to hibernating sites precludes the use of the term home range. Stomach contents of 21 G. geograph- ica, 54 G. ouachitensis and 35 G. pseudo- geographica females were quantitatively analyzed by both volume and frequency of occurrence. Graptemys geographica is a mollusk specialist, mollusks composed 66% of the volume of food consumed. Mollusks are also important in the diet of G. pseudogeographica females (19% by volume), but they make up only 2.8% of the diet of G. ouachitensis. Vegetation makes up a large proportion of the diet of G. ouachitensis (31.5%) and G. pseudo- geographica (42.4%). Insects composed 51% of the volume of food eaten by G. ouachitensis while only 21.9% and 15.3% in G. pseudogeographica and G. geo- graphica respectively. No differences in food habits were noted among the males of the three spe- cies. All consumed insect larvae, mol- lusks and fish carrion. Female G. ouachi- tensis were observed to be primarily sur- face feeders. Graptemys geographica and G. pseudogeographica feed mostly un- derwater. Map turtles become active in April and begin dispersing from the hi- bernacula while water temperatures are 4*^-7^C. During May females move out of the channels toward nesting beaches where they congregate to bask, but have not been observed feeding until 26 May; presumably they do not feed until after laying their first clutch of the year. In late May or early June map turtles lay their first clutch of eggs and then move away from the nesting islands to feed for two or three weeks while shells are being deposited around their second clutch of eggs. Feeding continues until 46 Tulane Studies in Zoology and Botany Vol. 22 the first week of September. By October all three species are moving toward com- munal overwintering sites. Up to 131 Graptemys have been caught in 30 mi- nutes behind a wing dam in October. Basking activity was observed primar- ily between 0900 and 1800 hrs. Through- out the day turtles alternate between basking, feeding or other activity in the water. Basking sites are used commun- ally. Up to 47 adult Graptemys were ob- served basking simultaneously on a 6 m log. No aggressive interactions were ob- served between basking Graptemys. On two different occasions a common grackle Quiscalus quiscula removed leeches {Placobdella parasitica) from basking Graptemys. Nest predation by red fox, raccoon and otter was observed. Hatchlings were con- sumed as they emerged from the nest by ring-billed gulls, crows, grackles and red- winged blackbirds. Outside of man, aduU map turtles have few potential predators in the study area. ACKNOWLEDGEMENTS This study was conducted under the supervision of Dr. William G. Reeder, who supplied field equipment, laboratory space, museum funds and technical criti- cism of the paper. Drs. Jack P. Hailman, C.J. McCoy, John Neess, M. Ewert and Ms. Marsha Christianson are thanked for their critical comments on the paper's composition. Dr. John Legler and Dr. Michael Ewert are appreciated for pro- viding me access to new techniques and data prior to publication. Cheryl Hughes is remembered for drawing all of the graphs. Mike Nee helped with the identification of plant species in the study area. Partial support for this study was provided by various travel grants: University of Wisconsin- Madison, Zoology Department Jefferson Davis Fund; University of Wisconsin- Madison Zoological Museum; Carnegie Museum of Natural History-Pittsburgh. Data for part of this study were collected while I was conducting research for the Wisconsin Department of Natural Re- sources, Bureau of Research. The DNR personnel were instrumental by lending of fyke nets and providing permits to con- duct my research. Throughout this study many students and friends gave freely of their time to help me with some aspect of the study - to all of them I express my sincerest grat- itude. Several of them gave up their sum- mers and worked for next to nothing both in the field and in the laboratory. Without their help this study would still be incom- plete. To each of them I offer my deepest appreciation: Marsha Christianson, George Dmytrenko, Janis I. Dzelzkalns, D.D. Flambeau Von Rex, Bruce Hell- mich, Jeremy Jacobs, Alan Jaslow, Kris- tine Klevickis, Lisa Mattick, Michael Pappas and Dan Peterson. Robert Peter- son made the laboratory part of this study possible. Without his contribution to the care of the hatchling turtles, monitoring of incubating eggs, care of the experi- mental turtles for seven years, and main- tenance of my laboratory facilities a re- duced study would have had to be under- taken. My gratitude is extended most sincere- ly to Jack Roberts for his assistance and guidance on the Mississippi River at Stoddard. He gave freely of his know- ledge of the river, saved turtles captured in conjunction with his commercial fish- ing operation and provided companion- ship and local history of the study area. Marsha Christianson provided much needed help in the field and laboratory and prevailed through many trying mo- ments in the editing of this paper. Her patience and understanding will long be remembered. I must also express my thanks to the other occupants of the Birge No. 1 Wisconsin Map Turtles 47 Hall Animal House for enduring floods and odors from the turtle growing room and my continual expansion. Carmelita Bell is thanked for typing the final version of the manuscript. LITERATURE CITED Atland, P.D. 1951. Observations on the structure of the reproductive organs of the box turtle. J. Morphol. 89:599-621. Auth, D.L. 1975. Behavioral ecology of basking in the yellow-bellied turtle, Chrysemys scripia scripta (Schoepff). Bull. Fl. State Mus., Biol. Sci. 20: 1-45. Bowler, J. K. 1977. Longevity of reptiles and amphi- bians in North American collections. S.S.A.R. Herp. Cir. #6: iv -¥ 32 pp. Boyer, D.R. 1965. Ecology of the basking habit in turtles. Ecology 46: 99-118. Bull, J. and R.C. Vogt. 1979. Temperature-depen- dent sex determination in turtles. Science 206: 1186-1188. Cagle, F.R. 1944. Home range, homing behavior and migration in turtles. Misc. Publ. Mus. Zool. Univ. Michigan 61: 1-34. 1946. The growth of the slider turtle, Pseu- demys scripta elegans . Amer. Midi. Nat. 36: 685- 729. 1950. The life history of the slider turtle, Pseudemys scripta troosti (Holbrook). Ecol. Monogr. 20:31-54. 1952. The status of turtles Graptemys pulch- ra Baur and Graptemys barboiiri Carr and Mar- chand, with notes on their natural history. Co- peia 1952: 223-234. 1953. Two new subspecies of Graptemys Chrysemys picta in southeastern Pennsylvania. Herpetologica 27: 135-141. and R. W. Barbour. 1972. Turtles of the Uni- pseudogeographica. Occ. Pap. Mus. Zool. Univ. Michigan 546: 1-17. Carr, A.F. 1949. The identity of Malacoclemmys kohnii Baur. Herpetologica 5: 9-10. Claflin, T. 1973. Operation and maintenance 9-foot navigation channel upper Mississippi River head of navigation to Guttenberg, Iowa. U.S. Army Corps, of Engineers St. Paul District XVII 4- 596 pp. and exhibit - XV -^ 273 pp. Christiansen, J.L. and E.O. Moll. 1973. Latitudinal reproductive variation within a single subspe- cies of painted turtle Chrysemys picta belli. Her- petologica 29: 152-163. Curtis, J.T. 1959. The vegetation of Wisconsin. Univ. Wis. Press Madison, Wis. xi -i- 657 pp. Dorando, S.L. 1979. A method to reduce error in weight estimation of freshwater turtles. Copeia 1979: 346. Ernst, C.H. 1971. Growth of the painted turtle. ted States. The University Press of Kentucky, Lexington, Ky. x + 347 pp. Ewert, M.A. and J.M. Legler. 1978. Hormonal in- duction of oviposition in turtles. Herpetologica 34:314-318. Gibbons, J.W. 1967. Variation in growth rates in three populations of the painted turtle Chrys- emys picta. Herpetologica 23: 303. 1968. Population structure and survivorship in the painted turtle, Chrysemys picta. Copeia 1968: 260-268. 1970. Sex ratios in turtles. Res. Popul. Ecol. 12: 252-254. Graham, T.E. 1971. Growth rates of the red-bellied turtle, Chrysemys rubriventris, at Plymouth, Mass. Copeia 1971: 353-356. Gray, J.E. 1831. Synopsis reptilium. Part 1. Cata- phracta. Tortoises, crocodiles and enaliosauri- ans. Treutal, Wurtz, and Co. , London. 78 pp. Hartweg, N. 1946. Confirmation of overwintering in painted turtle hatchlings. Copeia 1946: 255. Hildebrand, S.F. 1929. Review of experiments on artificial culture of diamond-back terrapin. Bull. U.S. Bur. Fish. 45: 25-70. Jackson, C.G. and J.D. Davis. 1972. A quantiative study of the courtship display of the red-eared turtle Chrysemys scripta elegans (Wied). Herpe- tologica 28: 58-64. Legler, J.M. 1960. The natural history of the ornate box turtle, Terrapene ornata ornata Agassiz. Univ. Kansas Publ. Mus. Nat. Hist. 11 (10): 527-669. 1977. Stomach flushing: a technique forche- lonian dietary studies. Herpetologica 33: 281- 284. Miller, M.R. 1959. The endocrine basis for repro- ductive adaptation in reptiles, p. 449-516. In A. Gorbman (ed). Comparative endocrinology, John Wiley and Son, N. Y. Moll, E.O. 1973. Latitudinal and intersubspecific variation in reproduction of the painted turtle, Chrysemys picta. Herpetologica 29: 307-318. and J.M. Legler. 1971. The life history of a neotropical slider turtle, Pseudemys scripta (Schoepff) in Panama. Bull. Los Angeles Co. Mus. Nat. Hist. Soc. (11): 102 pp. Newman, H.H. 1906. The habits of certain tor- toises. J. Comp. Neurol. Psychol. 16: 126-152. Patterson, R. and B. Brattstrom. 1972. Growth in captive Gopherus agassizi. Herpetologica 28: 169-171. Pieau, C. 1971. Sur la proportion sexuelle chez les embryons de deus Cheloniens (Testudo graeca 48 Tulane Studies in Zoology and Botany Vol. 22 L. et Emys orbicularis L. ) issue d'oefs incubes artificiellement. C.R. Acad. Sci. (Paris) 272: 3071-3074. 1972. Effets de la temperature sur le de- vel6ppement des glandes genitales chez les em- bryons de deux cheloniens, Emys orbicularis L. , et Testudo graeca L. C.R. Acad. Sci. (Paris) 274: 719-722. 1973. Nouvelles donnees ejperimentales concernant les effets de la temperature sur la differenciation sexuelle chez les embryons de cheloniens. C.R. Acad. Sci. (Paris). 277: 2789- 2792. Powell, C. 1967. Female sexual cycles of C/iryse/nys picta and Clemmys insculpta in Nova Scotia. Canadian Field Nat. 81: 134-140. Ream, C. 1967. Some aspects of the ecology of the painted turtles of Lake Mendota, Wisconsin. Ph.D. dissertation. Univ. of Wisconsin-Mad- ison xxiii -I- 375 pp. Risley, P.L. 1938. Seasonal changes in the testes of the musk turtle Sternotherus odoratus L. J. Mor- phol. 63: 301-317. Sexton, O.J. 1957. Notes concerning turtle hatch- lings. Copeia 1957: 229-230. 1959. Spatial and temporal movements of a population of the painted turtle, Chrysemys pic- ta marginata (Agassiz). Ecol. Monogr. 29: 113- 140. Shealy, R.M. 1976. The natural history of the map turtle, Graptemys pulchra Baur, in Alabama. Bull. Fl. State Mus. Biol. Sci. 21: 47-111. Smith, P.W. 1961. The amphibians and reptiles of IlHnois. Illinois Nat. Hist. Surv. Bull. 28: 1-298. Timkin, R.L. 1968. Graptemys pseudogeographica in the upper Missouri River of the northcentral United States. J. Herpetol. 1: 76-82. Vogt, R.C. 1978. Systematics and ecology of the false map turtle complex Graptemys pseudoge- ographica. Ph.D. dissertation. Univ. of Wiscon- sin-Madison xxiii -I- 375 pp. 1979. Cleaning/feeding symbiosis between Crackles (Quiscalus: Icteridae) and map turtles (Graptemys: Emydidae). Auk 96: 608-609. (in press) Systematics of the false map turtle complex Graptemys pseudogeographica. Bull. Carnegie Museum of Natural History. _. (in press) New methods for trapping aquatic turtles. Copeia 1980. Webb, R.G. 1961. Observations on the life histories of turtles (genus Pseudemys and Graptemys) in Lake Texoma, Oklahoma. Amer. Midi. Nat. 65: 192-214. Yntema, C.L. 1976. Effects of incubation tempera- ture on sexual differentiation in the turtle, Chely- dra serpentina. J. Morphol. 150: 453-462. May 26, 1980 A TAXONOMIC STUDY OF THE COOTER TURTLES, PSEUDEMYS FLORIDANA (LECONTE) AND PSEUDEMYS CONCINNA (LECONTE), IN THE LOWER RED RIVER, ATCHAFALAYA RIVER, AND MISSISSIPPI RIVER BASINS KENNETH MARK FAHEY* Department of Biology, The University of Southwestern Louisiana Lafayette, Louisiana 70504 ABSTRACT The taxonomy of the cooter turtles, Pseudemys floridana (LeConte) and Pseudemys concinna (Le- Conte) was studied in southwestern Louisiana to determine the relationship between these two tur- tles and to examine the validity of the taxonomic characters currently utilized. One hundred and sixty-two turtles were exam- ined from the lower Red River, the Atchafalaya River, and the Mississippi River basins. The color patterns and osteological characteristics were nu- merically scored. A discriminant analysis based on the characters of plastral pattern, carapace pattern, bridge markings, and the number of phalanges in the fifth toes of the hind feet, was conducted on these specimens. Two a priori groups of specimens with relatively "pure" characters were selected, one with P. floridana characters and one with P. con- cinna characters. From these a priori groups a set of discriminant coefficients was calculated for each character and all specimens were assigned Z-values based on these characters. A linear plot of the Z-values showed most speci- mens in the sample were intermediate, with Z-val- ues distributed between and overlapping both a pri- ori groups. The four characters used had little or no taxonomic value in separating these turtles. The wide range of Z-values found within individual clutches indicated interbreeding of specimens with widely varying characteristics. Correlation coeffi- cients for toe phalanx number and plastral pattern versus carapace length indicated these characters to be ontogenetic. The results show that Pseudemys floridana and Pseudemys concinna should be synonymized under the senior synonym, Pseudemys floridana, pending discernment of quantitative characters that will dis- tinguish/fondana and concinna as species. INTRODUCTION The North American emydid turtles of the genus Pseudemys have had a long, confused taxonomic history. Relation- ships among many of the species com- plexes are neither understood nor agreed upon. Pseudemys floridana (LeConte) and Pseudemys concinna (LeConte) are members of one such complex. A taxo- nomic study of these two species was con- ducted in the lower Red River, the Atch- afalaya River, and the Mississippi River basins in Louisiana (Fig. 1). The three subspecies reported to occur in this area according to Ernst and Barbour (1972), and Conant (1975) are Pseudemys flori- dana hoyi (Agassiz), Pseudemys concinna hieroglyphica (Holbrook), and Pseu- demys concinna mobilensis (Holbrook). The purpose of this study was to deter- mine the relationship between Pseude- mys floridana and Pseudemys concinna in * Present address: Department of Zoology- Entomology, Auburn University, Auburn, Alabama, 36830. EDITORIAL COMMITTEE FOR THIS PAPER: DR. JOHN W. CRENSHAW, JR. , Director, School of Biology, Georgia Institute of Technology, Atlanta, Georgia 30332 DR. FRANCIS L. ROSE, Professor of Biology, Texas University, Lubbock, Texas 79409 49 50 Tulane Studies in Zoology and Botany Vol. 22 southern Louisiana and to examine the validity of the taxonomic characters cur- rently being utilized to separate them. Pseudemys concinna ranges through- out much of the southeastern United States and contains the subspecies con- cinna, suwanniensis, mobilensis, texana and hieroglyphica. P. floridana occurs in approximately the same geographic range and contains the subspecies flori- dana, peninsularis, and hoyi (Crenshaw, 1955; Ernst and Barbour, 1972; Conant, 1975). According to Ernst and Barbour (1972) Pseudemys concinna is an inland turtle that inhabits mainly rivers, prefer- ring those with moderate currents, abun- dant aquatic vegetations, and rocky bot- toms. It occurs, however, in almost any aquatic habitat such as lakes, ponds, swamps, tidal marshes, oxbows, and ditches. P. floridana inhabits any aquatic habitat in the coastal plains, preferring those with slow currents, soft bottoms, and abundant aquatic plants. ARKANSAS TEXAS Figure 1. Study area which includes the Lower Red River, the Atchafalaya River, and the Missis- sippi River basins in Louisiana. Taxonomic history. — The species were first described by LeConte (1830) as Testudo floridana and Testudo concinna. The early taxonomic literature (Bona- parte, 1831; Gray, 1831, 1855, 1863; Du- meril and Bibron, 1835; Holbrook, 1836, 1838; Agassiz, 1857; Baur, 1893) in- volved primarily generic name changes and the addition of new subspecies. Carr (1935) considered Pseudemys floridana and P. concinna to be northern and southern representatives of the same species. In LeConte's original description P. floridana had page priority so he se- lected it as the name of the species. Carr (1937) synonymized P. mobilensis with P. floridana and named a new subspecies P.f. suwanniensis. Stejneger (1938) re- cognized a new subspecies, P.c. hoyi, for the specimens which Agassiz had called Ptychemys hoyi. Carr (1938) described P.f. peninsularis from Florida and included an analysis and key to the P. floridana group. He stated, "Due to inherent genetic instability, or to re-establishment of intercourse between previously isolated stocks, individual var- iation within a local population (even in a single litter) may result in phenotypes su- perficially more dissimilar than the actual races." He thought many descriptions stressed characters that were highly vari- able or sexually dimorphic and that be- cause of this the original descriptions were inadequate. He described the P. floridana group as " . . . a Rassenkreis which extends westward in two limbs from the Atlantic coastal floridana - an inland series {concinna, hieroglyphica, texana), and another in the coastal plain {peninsularis, suwanniensis, mobilen- sis).'' Carr (1952) redescribed the group and added P.f. hoyi. Crenshaw (1955), in his study of the Florida races P.f. floridana, P.f. peninsu- laris, and P.f. suwanniensis, stated that No. 1 Cooler Turtles 51 the relationships between all turtles of this complex were best shown by subdiv- iding the complex into two species, P. concinna and P- floridana. The species P. concinna included the subspecies, con- cinna, mobilensis, hieroglyphica, suwan- niensis, and texana. The species P. flor- idana included the subspecies floridana, peninsularis, and hoyi. The present status of these turtles is that suggested by Cren- shaw (1955). Many authors include these turtles in the genus Chrysemys (McDow- ell, 1964; Weaver and Rose, 1967), how- ever this has never been uniformly ac- cepted (Holman, 1977) and data present- ed by Vogt and McCoy (1979) suggest that they should be placed in the genus Pseudemys and separated from Chrys- emys. Locality records and status. - Speci- mens of P. floridana are not easily distin- guished from specimens of P. concinna on the basis of presently utilized taxo- nomic characters. Reported records for the three Louisiana subspecies show that many authors are unsure of the identity of specimens and unwiUing to commit them- selves as to the exact ranges for each sub- species. One aspect of the problem is the paucity of specimens. Another is the wide range of characters these turtles possess and the wide geographic range they oc- cupy. Three subspecies occur in Texas ac- cording to Brown (1950): P.f. hoyi, P.c. texana, and P.c. mobilensis. Brown refer- red to all as subspecies of Pseudemys flor- idana. He noted that the distribution of P.c. texana was unclear and that a num- ber of specimens from Bastrop County might prove to be intergrades with P.f hoyi. Webb (1970) listed Pseudemys f hoyi as occurring in eastern Oklahoma and remarked that P.c. heiroglyphica probab- ly also ranged into eastern Oklahoma where the two "occasionally hybridize." Two subspecies are reported from Ar- kansas, C.c. hieroglyphica and C.f hoyi (Conant, 1975). Michael Plummer (pers. comm. November 17, 1976) has advised me that P. concinna appears to predomi- nate in Arkansas populations but that most specimens appear to have interme- diate characters. The subspecies C.c. hieroglyphica and C.f. hoyi occur in the southern third of the state of Missouri where they hybrid- ize in nature (Anderson, 1965). Smith (1961) treated all Illinois speci- mens as hybrids of P.c. concinna x P.f. hoyi. His remarks included an opinion from Crenshaw which suggested that all southern Illinois turtles be regarded as hybrids because of introgression of P. floridana genes into the P. concinna pop- ulations of the lower Mississippi River Valley. Crenshaw believed this produced an intermediate form. Two subspecies, C.c. hieroglyphica and C.f. hoyi, occur in Kentucky (Bar- bour, 1971). The ranges of these turtles in Kentucky appear to be sympatric and Barbour commented that the two hybrid- ize freely with each other. Barbour (1971) and Conant (1975) showed C.c. hiero- gr^phica and C.f hoyi as occurring in ap- proximately the same ranges in Tennessee. Three subspecies of C. concinna occur in Alabama; they include C.c. concinna, C.c. hieroglyphica, and C.c. mobilensis (Conant, 1975). One subspecies of C. floridana, C.f floridana, occurs in Ala- bama (Conant, 1975; Mount, 1976). In dealing with the subspecies of P. concin- na. Mount (1976) stated that, "... the geographic variation was found to be in- consistent with previous reports and as- sumptions" and nearly all characters used to distinguish the subspecies had wide areas of overlap. He felt the designation of subspecies was largely arbitrary. He chose to allocate all P. concinna in Ala- bama to the subspecies P.c. concinna and 52 Tulane Studies in Zoology and Botany Vol. 22 noted that hybridization between P. con- cinna and P. floridana occurs frequently in the southeastern portion of the state. Existing records for these turtles in Louisiana (Viosca, 1923, 1926; Cagle and Chaney, 1950) do not give a good indica- tion of the species and subspecies which are found in various localities. Liner (1954) listed three Tulane specimens (TU-1104, 11046, 13618) as P. floridana. The first is from Vermilion Parish and the last two are from Terrebonne Parish. These specimens were examined and the data included in my statistical analysis of specimens. Keiser and Wilson (1969) reported three subspecies of these turtles, C.c. hie- roglyphica, C.c. mobilensis, and C.f. ho- yi, from Louisiana, Keiser (1976) referred to the two species as the Chrysemys flori- dana-concinna complex and used the pre- sence of a plastral pattern and the C- shaped mark on the second costal scutes of the carapace of C concinna to separate the species. He also mentioned possible skeletal characters separating them, but noted that few Atchafalaya Basin speci- mens fit the descriptions of the subspecies given by Carr (1952). The localities and specimens cited by Keiser (1976) are in- cluded in this study as they constitute the core of the data from which the present study grew. MATERIALS AND METHODS Specimens examined. — The specimens used in this study were collected primar- ily from the Atchafalaya River Basin from 1975 to 1977. Specimens from the Red River and Mississippi River basins were borrowed from the Tulane Univer- sity collection of amphibians and reptiles (TU). Atchafalaya River Basin speci- mens are from the University of South- western Louisiana collection of amphibi- ans and repitles (USL). A total of 162 specimens was rexamined. All specimens collected during the course of this study were catalogued into the USL collection (Appendix I). Method of collection. — Several types of traps (Legler, 1960) were used in the collection of aduh turtles. Fish, chicken, cottonseed cakes, lettuce, and waterme- lon were tried as bait. Although no adult P. floridana or P. concinna were caught, adults of P. scripta, and C. picta were caught in traps. Most observations of feeding suggest that P. floridana and P. concinna are primarily vegetarian as adults (Allen, 1938; Carr, 1952; Ernst and Barbour, 1972). The abundance of natural plant foods may account for my inability to trap adult specimens even when plant materials were used as bait. I obtained adult specimens by purchas- ing them from fishermen or by searching for nesting females along roads. Road collecting was most successful in the early morning from 5:00 to 8:00 A.M. (C.S.T.) and in late afternoon from 3:30 P.M. until dusk. Five adults from which I later ob- tained egg clutches were collected in this manner. A gill net was occasionally used in the collection of adult turtles, though with limited success. Two methods used without success for the collection of juvenile turtles were night collecting and snorkeling. The most successful method of obtaining small tur- tles was to dip-net basking animals from a canoe or powerboat. Collection methods used in other areas with clear water (Chaney and Smith, 1950) did not work well in central Louisiana. The waters fre- quented by these turtles in central Loui- siana usually are over 1 m deep and are often very turbid. All eggs were collected from gravid fe- males using Cagle's (1944) method. Spec- imens were pithed and the entire oviducts were removed. The eggs were removed from the oviducts and placed in an in- No. 1 Cooler Turtles 53 cubator. Five clutches were incubated us- ing Trotter's (1973) technique, with hatching success varying from 5% to 88*7^. Each clutch of hatchlings was kept ahve for several months for observation and to allow color patterns to become distinct. Osteological preparations. - Pha- langes of the fifth toes of the hind feet were examined by X-ray (adult speci- mens) or clearing and staining (hatchlings and juveniles). The clearing and staining procedure was, with modifications, that of Hardaway and Williams ( 1975). Description of taxonomic characters. - A number of taxonomic characters are presently used to differentiate subspecies of P. floridana and P. concinna in various parts of their ranges. They do not apply to all the subspecies and the reliability of each character varies depending on the subspecies and the locality. As many characters as possible were examined to determine which could possibly be used to separate P. floridana and P. concinna in the area studied. Two characters used in this study and referred to in most present taxonomic lit- erature (Carr, 1952; Crenshaw, 1955; Ernst and Barbour, 1972; Conant, 1975) are the plastral pattern and the carapace markinizs. P. concinna is reported to ha\e a C-shaped yellow line on the second costal B Figure 2. Pattern on the second costal scutes of the carapace; A. C-shaped pattern of Pseudemys concinna; B. Straight Une pattern of P. flori- dana. Figure 3. Typical plastral patterns; A. Pseudemys concinna; B. P. floridana. scute of the carapace (Fig. 2) and a plast- ral pattern that generally follows the seams of the plastral scutes (Fig. 3). P. floridana has either a straight line or an inverted Y-shaped line on the second cos- tal scute of the carapace (Fig. 2) and an mimacuiate plastron (Fig. 3). Both of these characters are presently used as the major taxonomic characters separating these species. Both are based on pigmen- tation which, if used alone, is not a reli- able taxonomic character at the species level. Variation in color and in the ar- rangement of pattern can result from non-genetic factors. Temperature may play an important role in pattern determi- nations especially during embryonic de- velopment (Fowler, 1970; Vinegar, 1973, 1974). Three characters that are used to dis- tinguish various subspecies of P. flori- dana and P. concinna are neck stripe pat- tern, jaw serration, and the pattern on the ventral surface of the marginal scutes. All specimens in my sample were scored for each of these characters. These char- acters are, however, probably too qual- itative and their range of variation too great to be useful in statistical analyses. The pattern of stripes on the dorsal surface of the neck is used to distinguish P.f. peninsularis from P.c. suwanniensis in Florida. P.f. peninsularis has a hairpin 54 Tulane Studies in Zoology and Botany Vol. 22 pattern while Pc. suwanniensis has a se- ries of straight lines (Fig. 4). Sample spec- imens were scored as continuously lined (C), discontinuously lined (D), or hairpin patterned (H), for right and left sides of the neck. Many specimens possessed dif- ferent combinations of these three cate- gories and some possessed intermediate pattern types. Jaw serration is used to distinguish P.c. texana from the other subspetfes of P. concinna and P. floridana. This character can also be used to distinguish three Flor- ida turtles, P.f. peninsularis, P.c. suwan- niensis, and P. nelsoni, from each other. The sample specimens were scored on the basis of whether their jaws were serrated (S) or unserrated (U) and whether the upper jaw was notched (N) or un-notched (no symbol). The range of variation in- cluded that which was found in the above mentioned species long with other varia- tions (Fig. 5). The pattern on the ventral surface of the marginal scutes is used to distinguish P. concinna subspecies from P. floridana subspecies in Florida. P. floridana re- portedly has few or no marginal markings while P. concinna may possess a variety of Figure 4. Neck stripe patterns; A. Pseudemys con- cinna suwanniensis ^B. P. floridana peninsularis. Figure 5. Jaw serrations found in various species and subspecies of Pseudemys; A. P. concinna con- cinna; B. P. floridana hoyi; C. P. concinna tex- ana; D. a Louisiana variation; E. P. nelsoni (also found in some Louisiana specimens). patterns (Fig. 6). Three predominant pat- terns occurred in the sample, but a fourth type, the absence of markings, was not observed. Patterns were scored as types one to four (Fig. 6). A character I have analyzed in detail is the number of phalanges of the fifth toes of the hind feet (Weaver and Rose, 1967). Associated with this is the fusion or separation of the astragalus and cal- caneum in the ankle (pers. comm. June 17, 1976, with unpublished manuscript attached from Francis L. Rose). These characters, when first used by Weaver and Rose (1967) in their study of the ge- nus Chrysemys, appeared to be different for P. floridana and P. concinna. P. flor- idana reportedly had two toe phalanges and a fused astragalus and calcaneum. P. concinna reportedly had three toe pha- langes and a separated astragulus and cal- caneum (Fig. 7). These differences were theorized to be related to the degree of terrestrial or aquatic habits of each spe- cies (unpublished manuscript from Fran- cis L. Rose). P. floridana, the more ter- restrial turtle, has a shorter fifth toe and a more solid foot for better support while walking on land. P. concinna has a longer fifth toe which allows increased webbing; the separate astragalus and calcaneum al- No. 1 Cooter Turtles 55 Figure 6. A. Locations of the pattern on the ventral surfaces of the marginal scutes. B. Pattern types found on the ventral surfaces of the marginals and the numerical scoring used to classify these. SO make its foot more flexible for swim- ming. Rose (pers. comm. October 11, 1976) has states that these characters may or may not be good taxonomic char- acters. Penial morphology of P. floridana and P. concinna was shown to be identical with that of P. scripta and P. nelsoni (Zug, 1966). Osteological characteristics of the skulls of emydid turtles were anal- yzed and used for taxonomic purposes by McDowell (1964). He drew a distinction between a rubriventris series and a flori- dana series but did not identify any differ- ences between P. floridana and P. con- cinna. Skull osteology and penial mor- phology were not examined in this study. The precopulatory behavior reported for P. floridana (Cagle, 1950) appears to be similar to that reported for P. con- cinna (Marchand, 1944; C.G. Jackson, 1972). If there are subtle behavioral dif- ferences between P. concinna and P. flor- idana they have not been reported. Statistical methods. — Markings from the following body and shell regions were recorded for each specimen: carapace, plastron, neck, bridge, and marginal scutes. Toe phalanx number on the fifth toes of the hind feet, fusion of the as- tragalus to calcaneum, jaw serration, and carapace length were also recorded. The color pattern data were qualitative and various systems of scoring patterns were devised for quantifying these data. The scoring systems used in this study were based on the key characters that have been used to separate P. floridana and P. concinna as species. Plastral pattern was quantified by counting the number of plastral scutes that contained dark pigmentation. A number ranging from 0 to 12 was assigned to each turtle on the basis of the number of scutes containing dark pigmentation (Fig. 8). Most hterature describes P. flor- idana as having an immaculate plastron, however, very few specimens in my sam- ple completely lacked dark plastral pig- mentation. I assumed individuals with low scores to be of P. floridana stock. Pattern on the second costal scutes of the carapace was more difficult to score. When a large sample of specimens was examined, a gradation from the straight- line pattern of P. floridana to the C-shape of P. concinna could be observed. Five numbered categories were defined and numerical scores were assigned as fol- lows: straight-line (1), branching (2), Y- shaped (3), X-shaped (4), and C-shaped Figure 7. Bones of the hind feet; A. Pseudemys concinna, showing three phalanges on the fifth toe and separate astragalus and calcaneum; B. P. floridana, showing two phalanges on the fifth toe and a fused astragalus and calcaneum. 56 Tulane Studies in Zoology and Botany Vol. 22 (5) (Fig. 9). The decision between an X- shaped and a C-shaped pattern was based upon whether the posterior branches touched the neural and marginal scutes (Fig. 9-4; X-shaped) or touched the third costal scute (Fig. 9-5; C-shaped). In some cases, especially in older specimens, this was difficult to accurately determine. If the branches were not complete a judg- ment as to their general direction had to be made. Right and left costal patterns were scored separately and later summed. Four scutes make up the bridge that c««iects the plastron to the carapace. Specimens were scored 0 through 4 for left and right bridges separately, accord- ing to the number of scutes with dark pigmentation (Fig. 10). Scores for right and left sides were later summed. Neck striping and marginal markings were too qualitative and variable to be used in the statistical analyses. Toe phalanx numbers on the fifth toes of the hind feet were recorded separately and later summed for use in the discri- minant analysis. The astragalus fused to the calcaneum was not used since these bones were separate in all but three spec- imens. A discriminant function based on four characters (plastral pattern, carapace pattern, bridge markings, and toe pha- lanx number) was computed using the 12 5 0 Figure 8. The scoring system for plastral patterns. Numbers indicate the number of plastral scutes containing dark pigmentation. kCi3 UiL^ Figure 9. Scoring system used for the second costal scutes of the carapace; (1) Straight-line; (2) Branching; (3) Y-shaped; (4) X-^shaped; (5) C- shaped. Three variations are shown for each category. No. 1 Cooter Turtles 57 Figure 10. Scoring system used for scutes making up the bridge. Pattern categories 1 through 4 are shown through a 0 category could also be found. BMD-04M program (Dixon, 1973). This program develops the linear function of the selected variables that gives the larg- est ratio of between-group variance to within-group variance. A series of discri- minant coefficients is calculated to give this ratio and thus maximize discrimina- tion between two a priori groups (Rao, 1952; Krishnaiah, 1966; Kendall, 1968; Gnanadesikan, 1977). In accordance with this program, two a priori groups of twenty individuals each were selected from the total sample of 162 specimens; one group of specimens exhibited "pure" P. floridana characters, and the other "pure" P. concinna charac- ters. Age and sex were not a basis for the selection of the a priori groups. From these a priori groups the set of discrimi- nant coefficients was calculated. The sum of these coefficients times their respec- tive character scores gave a value Z for each specimen. Using these Z-values, the program automatically classified into one group or the other all specimens not in- cluded in the a priori groups. By writing an addition to the program the Z-values themselves were obtained. Plotting indi- vidual Z-values on the discriminant axis showed which specimens in the unknown sample were within the range of either a priori group and which were intermediate (Rohwer and Kilgore, 1968; Thaeler, 1968;J.F. Jackson, 1973). In addition to the discriminant analysis, correlation coefficients among several characters were computed using the BMD-02D program (Dixon, 1973). This program computes simple correlation co- efficients. Early in this study I noticed that plastral pattern appeared to fade with age and that the number of toe pha- langes on the fifth toes of the hind feet seemed to increase from 2 to 3 with age. Correlation coefficients between these characters and carapace length were computed to determine if this relation- ship actually existed. RESULTS Statistical analyses. — The results of the discriminant analysis (Fig. 11 A) clearly show that, based on Z-values calculated from the four characters used, the sample does not fall into two distinct groups (species). There is also no large interme- diate group that would indicate only Fj hybridization. The sample is instead evenly distributed between, and over- laps, each a priori group. This indicates interbreeding of P. concinna and P- flor- idana, and backcrossing and interbreed- ing of the hybrids. 2 1.0 a. • - -—• ^: V-»-^ -I > o 1 D 1 J 1 F 1 M 1 A 1 M 1 J 1 J 1 A S 1 0 1 N 1 OB 07- 0.6- Q < 0 J o Q2 0.1 • 1974 (I), • IBI IdIjIfIm'aImIjIjIaIsIoInI MONTH Figure 2. Mean diameter of largest ova (lower graph) and mean percentage ovary weight (upper graph) for Notropis sabinae from Big Cow Creek, Texas (dashed Hnes), and Bayou Anacoco, Louisiana (solid lines), 1972-73 (no April sample), April 1974, and March-April 1976. Lines connect only data points for sequential samples within the same year. Numbers in parentheses indicate the number of observations on the mean when less than ten. Statistics of variability are available in Heins(1979). productive condition in early May 1973, (63% MA&PS, n = 30). Ova diameters and ovarian weights remained large from May through late September 1973 ( X = 0.57 - 0.67 mm, 3.8% - 5.9% body weight). In September, 77% (n = 26) of the females were reproductive. Ova diameters and ovarian weights de- creased to an average of 0.13 mm and 1.0% of body weight by mid-October, and almost all females were considered non-reproductive (98% IM&EM, n = 55). The reproductive periodicity of females in Big Cow Creek was similar to that in Bayou Anacoco (Fig. 2), but re- production ended somewhat earlier in 1973. Diameters of ova and ovarian weights decreased to 0.22 mm and 0.7% of body weight by late September, and only 2 of 32 females (6%) were con- sidered reproductive. The periodicity and variability in temperature, photo- period and stream flow accompanying these cycles are shown in Figure 3. After attaining reproductive condi- tion, individual females remained repro- ductive throughout the spawning season. Maturing ova were present in reproduc- ing females throughout the season. The ova development pattern of this species exhibits two complements of yolk- bearing ova, a stock of smaller maturing ova and a differentiated group of larger mature ova (Fig. 4). Such a bimodal dis- tributions is indicative of a variable series of spawnings with complements of ova released periodically over an extended reproductive season (Hickling and Rutenberg, 1936; Prabhu, 1956; Qasim and Qayyum, 1961; MacGregor, 1970). Reproductive periodicity in males: The cycles of testis and tubercle development were similar for both areas. Males taken in December and February had no tuber- cles although there were scars on a few; testes were small and cloudy to white in color. In March samples testicular activ- ity was indicated by enlargement of the testes; tubercles also developed in March. Testes and tubercles were gener- ally well developed from April into September; however, there was a de- cline of testis and particularly tubercle development in Big Cow Creek males in September, coincident with the decline of reproductive activity in females. Testes' were small in October and November. Very small degenerating tubercles were still present in October but were generally absent in November, excepting tubercle scars in some males. Reproductive effort and ovum size: There was a highly significant corre- lation between number of mature ova and standard length among females from both localities (Table 1). Regression lines expressing the relationship be- tween counts of mature ova and body size did not differ in slope (F = 0. 14; df = 1, 60; p > .05) or elevation (F = 2.20; df = No. 2 Life History o/Notropis 73 1 , 61 ; p > .05) when tested by analyses of covariance; therefore, the data from both locahties were combined. Counts of mature ova ranged from 1 13 to 423 for females 35.4 - 47.8 mm SL. The inter- cept, regression coefficient and correla- tion coefficient for the linear relation- ship between fecundity and standard length among females in the combined sample (Fig. 5, r^ = 0.57) are also given in Table 1. Log-log and semi-logarithmic transformations yielded similar r^ values, indicating that any of the three models adequately defined the relation- ship. However, logarithmic transforma- tion of both variables stabilizes the var- iance along the regression line (Bagenal, 1967) and is important for use in predic- tion; the resulting equation is (r^ = 0.53; F = fecundity, SL = standard length): log,, F = -2.8297 + 3.1953(log„,SL). There was a low but significant correla- tion between ovarian weight-somatic weight ratio and body size among females from Bayou Anacoco, but there was no significant correlation among females from Big Cow Creek (Table 1, Fig. 6). This notwithstanding, there was a low but highly significant correlation between ovarian weight-somatic weight ratio and body size among females in the combined samples. In general, these ratios ranged from 4.8 -15.2%. There was a low but significant corre- lation between mean mature ovum size ,-. 5.0 - CVJ O X 4.0 - 5 0) rr CO 3.0 - 2 o UJ 2.0 - X CD < 1.0 - -J >- o < (D Q Q 0 - 15 - t 10 - « « o~:::r-ig— — » I n I . I j'f'm'a'm'j'j'a's'o'n 30 - O ^ 25 UJ ct ^ 20 < Q. UJ 5 - H 10 - • •. ITT I u I . I M ' A ' M ' J MONTH A ' S ' O ' N Figure 3. Environmental variability and periodicity for Big Cow Creek, Texas and Bayou Anacoco Louisiana. Upper graph: mean monthly number of daylight hours (bold asterisks. Lake Charles LA) and mean monthly^scharge as cubic meters per second per square kilometer for Big Cow Creek (1952-76 op^n circles, soUd Une) and Bayou Anacoco (1969-73, solid circles, solid hne). Lo^^^/Pf " "^^^" "1°"'^'^ """ temperature for 1973 (dashed Une, DeRidder, LA), early 1974 (solid Ime), early 1976 (dotted hne). 74 Tulane Studies in Zoology and Botany Vol. 22 N = 478 >- o z bJ O LU cr < cr UJ -| 1 1 1 1 1 1 \ 1 I— 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 OVA DIAMETER (MM) Figure 4. Size frequency distribution for vitello- genic ova in a reproductive Notropis sabinae fe- male, indicating the typical pattern of ova matura- tion and recruitment. and body size among females from both localities (Table 1, Fig. 7), indicating a tendency for larger females to produce larger ova. Data are missing for one specimen fromBayou Anacoco; the ov- ary was not well preserved and many eggs broke when teased from the folli- cles. Analysis of covariance showed that regression lines fitted to the data for each locahty did not differ in slope (F = 0.53; df = 1, 59; p > .05) or elevation (F = 2.03; df = 1, 60; p > .05); therefore, data for both localities can be combined. The mean sizes of the mature ova measured ranged from 0.63 to 0.81 mm diameter and averaged 0.72 mm (S = 0.0452). Ad- dition of mean mature ovum diameter to the linear equation for fecundity did not significantly increase the i^ value (F = 2.260; df=l, 60; p> 0.05). Sex ratio, size at maturity and mean size of adult males and females: Chi-square tests indicated that there were no significant differences from a 1 : 1 sex ratio at either locality. The hetero- geneity X^ was not significant (p> 05) for either Big Cow Creek (0.7485, df =4) or Bayou Anacoco (5.3745, df =4); simi- lar results were obtained for the pooled X2 at the two respective stations (2.9823, df = 1; 1.6864, df = 1). The number of specimens originally examined from Big Cow Creek to de- termine size at sexual maturity was insuf- ficient; therefore, supplemental speci- mens taken in August 1973 and a portion of those taken in May 1977 were also examined. The supplemental specimens appeared to differ somewhat from those initially examined (May— July 1973) in that individuals seemed to mature at a 450 400 350 > 300 UJ CD i 250 200 - 150- 100- m * • * F=-5I6.I3I8+I79550(SL) 1 1 I r— 35 40 45 50 STANDARD LENGTH (MM) Figure 5. Relationship between number of ma- ture ova and standard length for Notropis sabinae females from Big Cow Creek (triangles), Texas, and Bayou Anacoco (dots), Louisiana, 1973-74 and 1976-77. No. 2 Life History o/Notropis 75 Table 1. Summary of linear regression analyses of ova number-body size, ovary weight-somatic weight ratio-body size, and ova diameter-body size relationships for samples of Notropis sabinae from Big Cow Creek, Texas, and Bayou Anacoco, Louisiana, 1973-1974 and 1976-1977; ns = non-significant difference, * = significant difference at .05 level, ** = significant difference at .01 level. Ova number-body size (SL) Locality n Intercept Regression Coefficient r Big Cow Anacoco COMBINED 24 40 64 -581.3956 -497.7474 -516.1318 19.2567 17.6721 17.9550 .772** .758** .753** Ovary weight- somatic weight ratio —body size (SL) Locality n Intercept Regression Coefficient r Big Cow Anacoco COMBINED 24 40 64 -3.3558 -4.3164 0.3003 0.3282 .362"^ .364* .368** Ova diameter-body size (SL) Locality n Intercept Regression Coefficient r Big Cow Anacoco COMBINED 24 39 63 0.4200 0.5136 0.4735 0.0075 0.0049 0.0060 .443* .377* .409** smaller size. Despite this complication the data are useful for a approximate comparison of size at sexual maturity in the two populations. Males matured at a smaller size than females at both lo- calities,completing maturation by 29 mm SL in Bayou Anacoco and 30 mm SL in Big Cow Creek. Females were not comp- letely mature until 32 mm in Bayou Anacoco and 33 mm in Big Cow Creek. The analysis of variance performed to test for significant differences in mean size of adult males and females from Big Cow Creek indicated a significant inter- action between the effects of sex and month (Table 2); therefore, a test of sim- ple effects was performed to determine when there were differences in mean size. This procedure indicated that females were significantly larger than males in July (p. < .05); all other differ- ences were non-significant. The F-value for interaction was non-significant for the Bayou Anacoco samples, as was the F-value for sex, indicating that there were no significant differences in the mean sizes of adult males and females. Significant differences between monthly 76 Tulane Studies in Zoology and Botany Vol. 22 samples were indicated for both locali- ties but were not analyzed. The results for the July sample from Big Cow Creek are unexplained; however, they may be due to sampling error. It seems reason- able to conclude that there were gener- ally no significant differences in the mean sizes of adult males and females at either locality. Growth and Population Age Structure Temperatures in the study area varied considerably over short periods (daily, weekly); but mean monthly temperature variation was moderate (Fig. 3). There- fore, annulus formation in this warm temperate region is quite variable. Further, annulus formation may be espe- cially weak for smaller individuals ex- periencing their first winter. Thus, scale analyses were combined with length fre- quency analyses to obtain the most reli- able age estimates. These analyses (Figs. 8, 9) indicate that the maximum age for 20.- 5 o < o CD >- o 15.- IQ- 5.- 0W= - 4.3164 t- 0.3282(SL) ^ I 1 I 30 35 40 45 STANDARD LENGTH (MM) 50 Figure 6. Relationship between ovary weight-somatic weight ratio and standard length for Noiropis sabinae females from Big Cow Creek (triangles), Texas, and Bayou Anacoco (dots), Louisiana, 1973-74 and 1976-77. .85 - .80 ^ .75 H < > o .70 - • •* * • • - —^^ • • « • - ^ • • • • • - • - • * * • .65 - • * - 0D = 0.4735 + 0.0060(SL) .60 - I I 35 40 45 STANDARD LENGTH (MM) 50 Figure 7. Relationship between mean size of unovulated, mature ova and standard length for Notropis sabinae females from Big Cow Creek (triangles), Texas, and Bayou Anacoco (dots), Louisiana, 1973-74 and 1976-77. most N. sabinae is about IV2 to 2 years. None of the scales examined from March or September 1973 Big Cow Creek sam- ples (Fig. 8; n = 50, 54, respectively) had more than two annual marks. Some specimens examined from March appeared to be forming an annulus. Th- ese data, and examinations of a few larger specimens in April and May col- lections (a time of active annulus forma- tion), suggest that some of the larger individuals (particularly those 44 mm SL) may enter a fourth growing season as three-year olds. Analysis of the length No. 2 Life History o/Notropis 77 c o (U U o U 00 c U o Q 1— I ON 3 1—1 OS cd CM (N X C/5 >o r^ 1-H ka 00 cd S m On ^ — ' On On' 00 ^-^ r-4 00 00 ^-^ 00 r*^ 00 ^-^ d ^ t/5 IT) t^ >^ rN -^^ Tf ^ "cd (N "^ ro o 1/5 (L) Id > I PU on on O 1/5 * O Q IT) (N Tt ^ O (N CO r- ^ (N in rsi r^ '— I m '*■ ^ d r~^ s ON ON 00 in r- 'i- 00 r*^ vO ^ ,^ (N 00 in ^ Tt 00 T^ rt 00 c o •5 2 •- c S Ci XI <^ ^ t ■4—* O 78 Tulane Studies in Zoology and Botany Vol. 22 CO m t^ Ov m m in ^^ 1— 1 /"-^ > O "^ (N 00 o • Tf • m l^ ^^ t^ -^ m m o o O en c < 3 O c o • ^^ ■«-' o "o U <*- o (U ■4-J cd Q ON a ON 3 I— > ON >-. CO On X CO On ON (N CO CM 00 ;-^ On "^ o 00 m 00 O m ^^ (^ 00 • 1— I 00 ^^ ON ^^ 00 CI 00 m (N E (/5 c c D o (4-1 o c < C/5 1/3 C/5 CJ 3 O C/5 c * c fN '-H (N O rj NO Tf NO r<^ ^" ^* o ^ (N l-H r^ ON in O 00 "^ IT) t^ 00 o lO r<-) On (N >o Tt ro NO ri NO <^ r-' 1— 1 00 (N (N ^ IT) rF <^ IT) 00 IT) NO m Tj- ON t^ NO t^ IT) 00 CN iri cn o 1— I -^ ^H m 1—1 CM Tj- Tt r- '■♦-< NO 00 «3 O H ^ O O 3 w > c 2 o c rs a* . C lU 3 2 id U J O ■- -^ OJ t/3 "" cd c o "^ « ^ ^ '-S °' c a xi ■<-' 3 o y E c/n N t/2 c cd O " C _ . o > ^ ^ ;n t/3 -3 No. 2 Life History o/Notropis 79 SEP AUG o z cr LlI LlI < 2 o tr JUL JUN MAY \- rO HIIUHimi III n N=58 N = 48 N>=38 N»I89 N-56 N = 85 jI-i.I ommm Itl N = 63 MAR I I 0) MAY APR MAR NOV OCT SEP N=230 N = II3 N = 85 N = 68 N = I2I N = I47 N = IIO T r" 11 II 1 T "T^ 1 20 30 40 5 0 0 10 STANDARD LENGTH (MM) 20 30 40 50 Figure 9. Length frequency histograms for collections of Notropis sabinae from Bayou Anacoco, Louisiana. Horizontal bars are placed at beginning of each month; histogram bases are set at date of collection. No. 2 Life History o/Notropis 81 frequency histograms for Big Cow Creek prior to and during the early part of the annual reproductive season demon- strates a bi-modal distribution. The ori- gins and fates of these two groups sug- gest that the larger group is primarily comprised of one-year old individuals entering their third growth season as two-year olds. The number in this group dwindles considerably by the end of this season, suggesting that most individuals die by the age of two years. The group of smaller fish thus represents individuals having just completed a first winter and entering a second growth season as one year olds. Maximum size was about 49 mm and 47 mm SL in Big Cow Creek and Bayou Anacoco, respectively. Finally, these data and the data on size at matur- ity suggests that a small percentage of individuals might mature within their first growth season. However, at least the majority mature in their second season, at or within one year. DISCUSSION Notropis sabinae, along with other members of the Notropis longirostris species group (Heins and Clemmer, 1976; Heins, 1979; Heins et al. , in press), has an extended reproductive season during which it seems a number of clutches of eggs are spawned. The long annual period of high temperatures and long photoperiods in the area (Fig. 3) probably facilitates the protracted repro- ductive season; temperature and photo- period are two proximal factors that acti- vate neuroendocrine centers controlling reproductive cycles in fishes (Schwass- man, 1971; DeVlaming, 1972, 1974). Nevertheless, extended reproduction may be adaptive to a variable environ- ment in that distributing reproductive activity over a long period of time may reduce the chance of losing a large por- tion of the annual recruitment (Starret, 1951; Tanyolac, 1973; Wallace,, 1973; Heins and Bresnick, 1975; Heins and Clemmer, 1976; Heins et al., in press). Protracted reproduction has also been observed in a number of southern spe- cies of Notropis (Mathur and Ramsey, 1974; Cowell and barnett, 1974; Beach' 1974; Cowell and Resico, 1975; Heins and Bresnick, 1975). The protracted reproductive season and presumed production of multiple clutches of eggs indicates a relatively high seasonal reproductive effort for Notropis sabinae. Further, N. sabinae is a relatively small, short-lived member of the genus Notropis as are other members of the A^. longirostris species group (cf. Hubbs and Hubbs, 1958; Carlander, 1969; Snelson and Jenkins, 1973; Hubbs and Miller, 1975; Heins and Clemmer, 1976; Heins, 1979; Heins et al. , in press). N. sabinae also matures early, at about one year. Williams (1966) suggested that a small, short-lived species with high an- nual mortality should invest a greater effort in reproduction in a given season, as compared to a large, long-lived species. Additionally, strong selection for early maturity, that is within one year, should result from a short life ex- pectancy (Tinkle, 1969). The spawning of N. sabinae was not observed during this study; however, it presumably occurs over open sand as in Notropis longirostris (Hubbs and Walker, 1942). If this is so, N. sabinae would belong to the ecological guild of psammophilous fishes delineated by Ba- lon (1975), as do other members of the N. longirostris species group. Notropis sabinae, as with other mem- bers of the N. longirostris species group, exhibits life history traits conventionally considered characteristic of a relatively r-selected species (cf. Steams, 1976; Ba- lon et al., 1977). Nevertheless, some in- terspecific variation was observed among the life history patterns of mem- bers of the N. longirostris species group; variation was also noted among papula- tions of yV. longirostris which was more widely studied (Heins, 1979). This intra- and inter-specific life history variation will be considered in a forthcoming re- port. However, I will consider a distinc- 82 Tulane Studies in Zoology and Botany Vol. 22 live difference between N. sabinae and other members of the N. longirostris species group herein. Among those populations studied thus far, N. sabinae produces smaller mature, unovulated ova (cf. Heins and Clemmer, 1976; Heins, 1979; Heins et al., in press). The inter- and intra-specific variation observed in egg size appears to be re- lated,at least in part, to stream flow pat- terns in southern North America (Fig. 10). Larger egg sizes were found in popu- lations inhabiting areas of higher annual stream runoff (Heins, 1979). I postulate that there has been a selective advantage to larger egg size in those streams where average annual runoff is greater. Thus annual stream runoff values may indi- cate the relative severity of this factor in the respective stream environments. Svardson (1949) and Williams (1959) have argued that a reduction in fecundity could not be favored by natural selection unless it was a necessary consequence of some advantageous development such as increased egg size. Larger ova generally result in larger larvae that can be expected to be stronger, able to swim better, and less susceptible to damage (Blaxter, 1969). Thus, larger ova may SCALE OF MILES 0 500 Figure 10. Map showing locations of various populations of Notropis sabinae (triangles), Notropis longirostris (dots), and Notropis species (cf. N. longirostris) (star) studied by Heins (1979) in relation to amount of annual stream runoff (cm). (After Geraghty et al. ( 1973), by permission. ) result in larger larvae better able to sur- vive in streams with greater average an- nual runoff. Heins and Clemmer (1976) had intimated this in their report on N. longirostris. This notion will be treated more fully along with other inter- and intra-specific variations in the life histories of members of the Notropis longirostris species group in a forthcom- ing paper. ACKNOWLEDGMENTS Field sampling would have been im- possible without the continued assis- tance of David Sever and Mike Sobczak. Gerry Bresnick, Bob Cashner, Art Johnson/ Dave Myers, Pat Sorenson and Steve Stevenson also assisted with the sampling. Gerri Rehbein helped with some early laboratory analyses. My ma- jor professor, Gerald E. Gunnmg, has supported my research in many im- portant ways and reviewed the original manuscript. William Dunlap aided in some statistical analyses. Mr. and Mrs. Ben P. Wilson of Call, Texas, graciously allowed me access to Big Cow Creek from their property and are deserving of special thanks. Eugene C.Beckham, III prepared the drawing of head tubercula- tion. Rebecca Gardner typed the manus- cript. This study was supported by a Sigma Xi Grant-in-Aid of Research. LITERATLfRE CITED BAGENAL, T.B. 1%7. A short review of fish fecundity, p. 89-1 1 1 . In: The biological basis of freshwater fish production. Gerking. S.D. (ed), Blackwell Scientific Publ, Oxford. BALON, E. K. 1975. Reproductive guilds of fishes: a proposal and a definition. J. Fish. Res. Board Can. 32:821-864. W. T. MOMOT and H. A. REGIER. 1977. Reproductive guilds of per- cids: results of the paleogeographical history and ecological succesion. J. Fish. Res. Board Can. .14: 1910-1921. BEACH, M. L. 1974. Food habits and reproduc- tion of the taillight shiner, Notropis maculatus (Hay), in central Florida. Florida Sci. 37: 5-16. BLAXTER, J. H. S. 1%9. Development: eggs and larvae, p. 177-252. In: Fish physiology. Vol. 3. Hoar, W. S. and D. J. Randall (eds.). Aca- demic Press, New York. No. 2 Life History o/Notropis 83 BUCHANAN, T.M. 1973. Key to the fishes of Arkansas. Arkansas Game and Fish Commis- sion, Little Rock, Arkansas. CARLANDER, K. D. 1969. Handbook of fresh- water fishery biology. Vol. 1. Iowa State Uni- versity Press, Ames, Iowa. COWELL, B. C. and B. S. BARNETT. 1974. Life history of the taillight shiner, Notropis macu- latus, in central Florida. Amer. Midi. Nat. 91: 282-293. , and C. H. RESICO, JR. 1975. Life history patterns in the coastal shiner, Notropis petersoni. Fowler. Florida Sci. 38: 113-121. DeVLAMING, V. L. 1972. Environmental con- trol of teleost reproductive cycles: a brief re- view. J. Fish Biol. 4: 131-140. .1974. Environmental and endocrine control of teleost reproduction, p. 13-83. In: Control of sex in fishes. Schreck, C.B. (ed.). Extension Division, Virginia Polytechnic Inst, and State Univ. DOUGLAS, N. H. 1974. Freshwater fishes of Louisiana. Claitor's Publishing Division, Baton Rouge, La. GERAGHTY, J. J., D. W. MILLER, F. V. D. LEEDEN and F. L. TROISE. 1973. Water Atlas of the United States. Water Information Venter, Port Washington, NY. HEINS, D. C. 1979. A comparative life history of a closely related group of minnows (Notropis, Cyprinidae) inhabiting streams of the Gulf Coastal Plain. Unpublished Ph.D. Disserta- tion, Tulane Univ., New Orleans, LA. , and G. I. BRESNICK. 1975. The ecological life history of the cherryfin shiner, Notropis roseipinnis. Trans. Amer.Fish. Soc. 104:516-523. and G. H. CLEMMER. 1975. Ecol- , and B. W. WALKER. 1942. Habi- tat and breeding behavior of the American cyprinid fish Notropis longirostris. Copeia 1942: 101-104. 1968. Fishes, p. 21-165. 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C. 1970. A review of the eastern North American cyprinid fishes of the Notropis tex- anus species group (subgenus /t/fcurnopi), with a definition of the subgenus Hydrophlox, and 84 Tulane Studies in Zoology and Botany Vol. 22 materials for a revision of the subgenus Alburnops. Unpublished Ph.D. Dissertation, Florida State Univ., Tallahassee, FL. TANYOLAC, J. 1973. Morphometric variation and life history of the cyprinid fish Notropis stramineus (Cope). Occ. Pap. Mus. Nat. Hist., Univ. Kansas 12: 1-28. TINKLE, D. W. 1969. The concept of reproduc- tive effort and its relation to the evolution of Ufe histories of lizards. Amer. Nat. 103: 501-516. WALLACE, D. C. 1973. Reproduction of the silverjaw minnow, Ericymba buccata Cope. Trans. Amer. Fish. Soc. 102: 786-793. WILLIAMS, G. C. 1959. Ovary weights of darters: a test of the alleged association of pa- rental care with reduced fecundity in fishes. Copeia 1959: 18-24. . 1966. Natural selection, the costs of reproduction, and a refinement of Lack's principle. Amer. Nat. 100: 687-690. THE SYSTEMATICS AND DISTRIBUTION OF THE HOGNOSE VIPER BOTHROPS NASUTA' BOCOURT (SERPENTES: VIPERIDAE) LOUIS PORRAS 12120 S.W. 180th Street, Miami, Florida 33177 JAMES R. McCRANIE 10770 S.W. 164th Street, Miami, Florida 33157 and LARRY DAVID WILSON Division of Intercurricular Studies, Miami- Dade Community College, South Campus, Miami, Florida 33176 ABSTRACT Variation in selected characters of scutellation, coloration, and body proportions in the Neotropi- cal hognose viper, Bothrops nasuta Bocourt, is dis- cussed and the specied is redescribed. Analysis of this variation allows for the definition of three dis- tinctive populations. No infraspecific taxa are rec- ognized. The biogeographic history of the species is also discussed. INTRODUCTION Bothrops nasuta Bocourt, 1868, is a small, terrestrial pit viper occurring in humid lowland forests from Chiapas, Mexico southward to Ecuador. It is a member of the Bothrops lansbergi group, a group of hognose vipers {sensu Burger, 1971), which was last revised by Amaral (1929a). The hognose viper group is in need of systematic study and the acquisition of an unusual specimen of B. nasuta from Ecuador by one of us (LP) prompted a review of the systematics of this species, the most widespread mem- ber of the group. Amaral (1929a) had 32 specimens a- vailable to him. Since his revision more material has become available allowing for a more complete understanding of the systematics of this species. MATERIAL AND METHODS We examined 241 specimens of Bothrops nasuta. The single specimen of this species from Belize (Neill, 1%5) was not available to us. The following scales were counted using the definition of Klauber (1972): intercanthals, prefove- als, preoculars, suboculars, and postocu- lars. We follow Burger (1971), with some modification, in defining and re- cording the following scales: post- canthals, interictals, interoculars, ocula- bials, and nasorostrals. The definitions of these latter scales are as follows: Postcanthals. The undifferentiated marginal head scale(s) contacting the upper preocular between the canthal and supraocular. Interictals. The supracephalic scales occupying the posterior portion of the head from oral rictus to oral rictus. For consistency, the interic- tals were counted between the last supralabials. Interoculars. The irregularly disposed scales in the frontal region that were counted along the path be- tween the supraoculars that trav- erses the fewest scales. Oculabials. The rows of scales between 'According to Shreve (1957), Amaral (1964a, 1964b, 1976) and Schwartz and Thomas (1975), generic names ending in -ops are feminine in classical Greek usage. Thus, the specific name for this hognose viper is nasuta, not nasutus. For opinions to the contrary, seethe 1964 edition of the International Code of Zoological Nomenclature and Smith and Larsen (1974). EDITORIAL COMMITTEE FOR THIS PAPER: DR. JONATHON A. CAMPBELL, Curator of Reptiles and Amphibians, Uni- versity of Texas at Arlington, Arlington, Texas 76019 DR. ROBERT A. THOMAS, Director, Louisiana N^^re Center, New Orieans, Louisiana 70127 85 86 Tulane Studies in Zoology and Botany Vol. 22 suboculars and supralabials but in- cluding neither. Nasorostrals. The scales between the nasal and rostral. The number of ventral scales was counted using the method of Dowling (1951). Due to the irregularity of scutel- lation in the nasofrontal area, the scales anterior to and including the interocu- lars, except for those bordering the edge of the head (rostral, intemasals, can- thals, postcanthals, and supraoculars), were counted and given the name nasofrontal. Postfoveals and median gulars were not counted due to their high degree of variability and consequent dif- ficulty of accurate tabulation. The dorsal scale rows were counted one head length posterior to the interictals, at midbody, and at the vent. All measurements were made on preserved specimens, with the exception of those made on a record- length specimen from Ecuador in the senior author's live collection. Statistical analyses were accomplished using the Student's t test, as described by Simpson, et al. (1960), using a probabil- ity value of .95. Inasmuch as Bothrops nasuta has a basically linear distribution, we used, for the sake of convenience, the political boundaries within the animal's range as the basis for subdividing for analysis our data on variation. The populational groupings in the tables are defined in the discussion. HISTORICAL SUMMARY A considerable amount of confusion has existed in the literature concerning the hognose vipers. Much of the confu- sion was largely eliminated by Amaral's paper (1929a) on the Bothrops lansbergi group. In that paper Amaral redescribed Bothrops nasuta Bocourt and demon- strated its distinctness from B. lansbergi (Schlegel). He also snyonymized B. brachy stoma (Cope) with B. lansbergi, thus terminating the application of the name brachystoma to specimens of B. nasuta. Moore (1962), however, incor- rectly placed brachystoma as a synonym of B. nasuta. Cope (1876) described Bothriopsis proboscideus. Later, he (1879) ques- tioned the validity of B. proboscideus but continued to recognize it in a later paper (Cope, 1887). Amaral (1929a) placed B. proboscideus in the synonymy of Bothrops nasuta. Posada- Arango (1889a) proposed a new genus of solenoglyph snakes, Thanatos, and described several new species, including T. sutus. Later in the same year (1889b) he introduced the generic name Thanatophis, without ex- planation (fide Vanzolini, 1977). Niceforo Maria (1938) concluded that Thana- tophis sutus is probably a synonym of Bothrops nasuta. His decision was tenta- tive due to Posada- Arango's generalized description. Garcia (1896), using the nomenclature of Posada-Arango, fig- ured a specimen identified as Thanatophis sutus, which is actually a juvenile B. nasuta, supporting the con- tention that the two names are synonym- ous, a supposition supported by Daniel (1949, 1955), Burger (1971), and us. Pe- ters and Orejas-Miranda (1970) and Van- zolini (1977) incorrectly place Thana- tophis (or Thanatos) sutus as a synonym of Bothrops lansbergi. Burger (1971) subdivided the various members of the genus Bothrops (sensu lato) into five genera. Bothrops nasuta was placed in the genus Porthidium as P. nasutum. This disposition was accepted by Smith and Smith (1976). Whereas we are in fundamental agreement with the generic limits proposed by Burger (1971), we prefer not to use them inasmuch as an adequate analysis of the basis for such a classification was not presented by him. In addition. Burger (1971) recognized two subspecies, nasu- tum and sutum {= nasuta and suta). This disposition was unsupported with any data and is not followed by us (see analysis of variation). No. 2 Hognose Viper 87 SPECIES ACCOUNT Bothrops nasuta Bocourt Bothrops lansbergii: Gimiher, 1863:350; Dunn, 1928: 30 (part); Picado.1936: 392; E. H. Taylor, 1951: 178, 1954: 681. Bothrops nasutus Bocourt, 1868: 202 (type locality: Panzos, on the banks of the Rio Polochic, Guatemala; holo- type: MNHN 1592 - fide Stuart, 1963), 1876: 410; Barbour and Loveridge, 1929: 3; Smith, 1943: 400, 1958: 224; Smith and Taylor, 1945 182; Stuart, 1948: 87, 1950: 24, 1958 29, 1963: 130; E. H. Taylor, 1954 783; Moore, 1962: 62; Duellman 1963: 245, 1966: 705; Klemmer, 1963 408; Neill, 1965: 113; Hoge, 1965 127; Savage, 1966: 751, 1973: 17 Heyer, 1967: 267; Moore, et al., 1968 48, 61; Meyer, 1969: 415; Minton and Minton, 1969: 207; Scott, 1969: 232 Peters and Orejas-Miranda, 1970: 49 Hoge and Romano, 1971: 252 Alvarez del Toro, 1973: 158; R. T Taylor et al., 1974: 384, 388 Henderson and Hoevers, 1975: 51 Campbell, 1976: 153. Porthidium nasutus: Cope, 1871: 207. Porthidium nasutum: Cope, 1871: 207, 1876: 151, 1879: 271, 1887: 89; Smith and Smith, 1976: S-B-16. Bothriopsis proboscideus Cope, 1876: 150 (type locality: Sipurio, at base of mountains, Costa Rica; no holotype designated); Cope, 1879: 271. Bothrops lansbergi: MuUer, 1878: 703 (in error). Bothrops brachy stoma: MuUer, 1882: 154; Amaral, 1925: 29, 1927a: 22, 1927b: 47; Loveridge, 1928: 63. Bothriopsis (sic) proboscideus: Cope, 1887: 89. Brothriopsis brachystoma: Cope, 1887: 89 (part). Thanatos sutus Posada- Arango, 1889a: 45-49 (paper not seen) . Thanatophis sutus Posada- Arango, 1889b: 344 (type locality: "le district de Zea," Colombia; no holotype de- signated); Garcia, 1896: 26. Bothriechis lansbergii: Gunther, 1895: 190 (part). Lachesis brachystoma: Boulenger, 1896: 547 (part); Boettger, 1898: 139. Trimeresurus brachystoma: Mocquard, 1909, 945 (part). Bothrops nasuta: Amaral, 1929a: 25. 26, 1929b: 237, 1931a: 89, 1931b: 94, 1935: 22, 1944a: 7, 10, 1944b: 18; Picado, 1931a: 69, 1931b: 104, 1936: 391; Dunn and Emlen, 1932: 32; Dunn, 1933: 79, 1944: 215; Schmidt, 1933: 19; Wettstein, 1934: 38; Nice- foro Maria, 1938: 419, 1942: 101; Prado, 1939: 1; Rendahl and Vestergren, 1941: 16, Daniel, 1949 329, 1955: 80; Pifano and Romer 1949: 302; E. H. Taylor, 1951: 177 Peters, 1960: 510; Villa, 1962: 50 Brattstrom 1964: 189; Medem, 1968 194; Minton and Minton, 1969: 216 Wilson and Meyer, in press. Trimeresurus nasutus: Dunn and Bailey, 1939: 20; Smith, 1941:62. Porthidium nasutum nasutum: Burger, 1971:35,132. Porthidium nasutum sutum: Burger, 1971: 35, 132. Description - The scutellation of this species is as follows: rostral higher than wide, the ventrolateral portion infre- quently separated off as a nasorostral (5.0% of specimens examined); inter- nasals paired, elongate, elevated anter- iorly, usuaUy in contact (81.4%); can- thals usually single (%.6%); postcanthals often one (67.7%), sometimes two; m- tercanthals 3-7 (x=4.95); nasofrontals 20-59 (x= 37.23); interoculars 3-7 (x=5.19); lateral edge of supraoculars flattened; interictals 19-28 (x=24.58); nasal" scale partially divided; prefoveals 2-12 (x=5.60); subfoveals 1-7 (x=2.26); loreal wider than high, upper preocular large, extending dorsally over the can- thai ridge; middle preocular divided or not, infrequently absent, in contact with orbit or not; lower preocular single, in- 88 Tulane Studies in Zoology and Botany Vol. 22 frequently absent, in contact with orbit or not; suboculars 1-4 (x=1.66); postoculars 1-4 (x=2.30); oculabials 1-4, usually 2 (88.8%); supralabials 8-11 (x=9.34); infralabials 9-13 (x= 11.41), the first pair usually in contact; a single pair of chinshields; ventrals 123-145 (x= 135.80); anal plate single; subcaud- als entire, 24-41 (x=30.02); tail not pre- hensile; dorsal scale rows at neck 23-29, usually 25 (87.8%), at midbody 21-27, usually 23 (92.8%), at vent 17-21, usu- ally 19 (96.6%); apical pits absent. The hemipenis is divided with a bi- furcate sulcus spermaticus, the sulcus di- viding close to the base of the organ and each branch extending to the distal end of the apical lobe. The basal portion of the organ and the areas lateral to and between the branches of the sulcus are spinulate. The shoulders and the absul- cate side of the organ are covered with spines which increase gradually in size proximally, terminating with a pair of enlarged basal spines situated on either side of the sulcus. Distal areas of the apical lobes are calyculate with papillate micro-ornamentation. Distribution and Ecology - Bothrops nasuta is an inhabitant of the humid low- lands of Middle America and adjacent northwestern South America (Figs. 1 and 2). Bothrops nasuta belongs to the Eatem Mesoamerican Complex of Savage (1966) and the Humid Tropical Assemblage of Duellman (1966). It is distributed along the Caribbean low- lands from northern Chiapas, Mexico, eastward through northern Guatemala, possibly to Belize (see discussion be- low), thence southward through Central America (Fig. 1) and onto the Pacific coastal plain of western Columbia to midwestem Ecuador (Fig. 2). The species probably also occurs in the mesic lowlands of eastern Veracruz, Mexico, inasmuch as it has been recorded near the border in Chiapas in the same type of forest (Alvarez del Toro, 1973). A speci- men (ANSP 4873) recorded from "Vera- cruz, Mexico" shows characters of speci- mens from much farther south in the range and we regard the data as ques- tionable. The species probably also occurs in the southern portions of Tabasco. The occurrence of Bothrops nasuta in Belize is questionable. Schmidt (1941) recorded a specimen of a hognose viper from Benque Viejo, Cayo District, Be- lize, which he identified as Trimeresurus yucatanicus. Neill (1965) recorded a specimen of B. nasuta from Xuan- tunich ( = Benque Viejo), Cayo District, and speculated that Schmidt's specimen was probably of the latter species. This disposition was followed by Henderson and Hoevers (1975). McCranie and Por- ras (1978) examined the Schmidt speci- men (USNM 61781), however, and de- termined it to be Bothrops yucatanica, thus reestablishing the occurrence of this species in Belize. The description of the specimen reported by Neill (1965) is am- biguous and will not distinguish B. nasuta from B. yucatanica. In the northern part of its range (the Mexican and most of the Guatemalan portion), B. nasuta lives in the broad- leafed forest or quasi-rainforest division of the evergreen forest (c.f. Duellman, 1966; also see Stuart, 1966). Duellman ( 1966) characterized these forests as hav- ing a marked dry season at which time some of the trees become leafless. How- ever, some rain apparently falls through- out the year (Duellman, 1963). Stuart (1958) considered the forests of El Pe- ten, Guatemala, as representing a transi- tion from the wet forests of the south and the dry forests of the outer Yucatan Peninsula to the north. Bothrops nasuta apparently adapts to the drier conditions of the quasi-rainforest by living in thick woods around marshy areas (Alvarez del Toro, 1973), in the vicinity of rivers where the forests are more mesic (Chama, 12 km NW Chinaja, Panzos, Piedras Negras, and Sayaxche, Guate- mala), under similar conditions around Lago Miramar, Chiapas, Mexico, and in the high forests surrounding the aguadas No. 2 Hognose Viper 89 of the Peten region of Guatemala. From extreme eastern Guatemala southward throughout Central America, the range of B. nasuta is within the more humid tropical rainforest division of the everg- reen forest (c.f. Duellman, 1966). This forest differs from the quasi-rainforest in that the habitat is at least moderately moist throughout the year even though rainy and dry seasons are evident. These moist conditions are enhanced by the tendency of the forest to develop a continuous treetop canopy which provides abundant shade creating the "greenhouse effect" discussed by Duellman (1966). The only known occurrence of B. nasuta outside the evergreen forest is in the scrub forest (c.f. Duellman, 1966) of the Sula Plain near San Pedro Sula, Honduras. How- ever, we believe these specimens came from gallery forest associated with the rivers of the region. In several places in Costa Rica and Panama, Bothrops nasuta crosses the Continental Divide onto the Pacific ver- sant. The species has been collected on both sides of the divide in the Tilaran area of Guanacaste Province, Costa Rica. Along the Tilaran transect studied by heyer (1967), the range of fi. nasuta is apparently continuous across the low di- vide (850 m) onto the Pacific versant, where the species has been collected ap- proximately 4 km west of the divide. One Figure 1 . Distribution of Bothrops nasuta in Central America. Solid symbols represent localities of specimens examined. Hollow circles represent literature records (see section on distribution and ecology for discussion of questioned locality in Belize). The type locality is indicated by a star within a circle. 90 Tulane Studies in Zoology and Botany Vol. 22 specimen (UCR 110) was collected on the Pacific versant at Puriscal (= Santiago), San Jose Province, Costa Rica. This specimen represents a relict population. An isolated population also occurs on the Pacific lowlands of the Golfo Dulce area of Costa Rica and probably adjacent Panama. Savage (1966) and Savage and Vial (1974) dis- cussed the factors creating the isolation of the vegetation of this region. How- 80" Figure 2. Distribution of Bothrops nasuta in Panana, Colombia, and Ecuador. For explanation of symbols see Figure 1. ever, the herpetofauna of the Golfo Dulce area resembles that of the Carib- bean lowlands of Costa Rica and Panama (Duelhnan, 1966; Savage, 1966; Savage and Vial, 1974). In Panama, B. nasuta has been collected on the Pacific versant close to the Continental Divide in El Valle de Anton, Code Province. Dunn (1933), in a discussion of the El Valle area, stated that "Atlantic side conditions and fauna come a little way over the divide onto the Pacific versant." Two specimens collected by Herbert Clark (MCZ 37115-16) are recorded from Panama City, Panama. Panama City is located in a savanna habitat (c.f. Duelhnan, 1966). These two specimens probably came from farther north in the mesic forests of the Canal Zone area where Clark obtained 13 other speci- mens. No specimens are available from Panama east of the Canal Zone area, but it undoubtedly occurs in the Caribbean mesic forests of eastern Panama'. In South America Bothrops nasuta in- habits the lowland rainforests of Colom- bia west of the Andes southward to midwestem Ecuador. Dunn (1940), Duelhnan (1966), and Savage (1966) dis- cussed the "crossover effect" occurring in the eastern Panama-northwestern Colombia region, where the Caribbean mesic forms of lower Central America cross onto the wet Pacific lowlands of Colombia. However, the Pacific low- lands consist of a very narrow strip in northwestern Colombia. The lowland rainforests east of the Andes in the cen- tral jxjrtion of the department of Choco and the adjacent western part of the de- partment of Antioquia, Colombia, which are also inhabited by B. nasuta, are drained by the Rio Atrato and its tributaries, which then flow northward into the Caribbean Sea. It is not until the Ri'o San Juan drainage of southern Choco is reached that B. nasuta becomes essentially a Pacific coastal plain animal, which it then remains to the terminus of its range in midwestem Ecuador. Bothrops nasuta also inhabits the low- No. 2 Hognose Viper 91 land rainforests of the rios Cauca and Force in the department of Antioquia, Colombia. As with the Rio Atrato, these latter two rivers flow northward into the Caribbean. The forests along the eastern edge of the Andes and on the Pacific coastal plain of Colombia are one of the most humid areas of the world and certainly the most humid of all of South America (Eidt, 1968; Haffer, 1970). The rains in the Quibdo area, Choco, Colom- bia, fall year around (Haffer, 1970). Eidt (1968), in describing central Choco, Col- ombia, stated that "only for a few days each year, usually when winds from the north disturb the general calm of the re- gion, are there brief respites from the near constant rains." Myers et al-(1978) discussed the habitat at Quebrada Guangui, Depto. Cauca, Colombia, where they collected a series of B. nasuta (AMNH 109794-811). Bothrops nasuta occurs from near sea level to about 900 m in elevation, but is generally found below the 600 m con- tour. Of 112 specimens with elevational data, 49 (43.8%) were from below 200 n and 107 (95.5%) were from below 625 m The highest elevation datum available i 900 m for two specimens (UMMi 116523, 117735) from near Matagalpa Nicaragua. Dunn (1944) gave an eleva tion of 1885 m for B. nasuta based on a record for Urrao, Antioquia, Colombia (Niceforo Maria, 1938, 1942). We doubt that the specimen actually came from this elevation. Niceforo Maria (1938) stated "que le fueron remitidos de. . .." This literally means "received from," which we interpret to mean "shipped from." We also question two other Niceforo Maria (1938, 1942) records; Pueblo Rico, Depto. Caldas and Medel- li'n, Depto. Antioquia. The elevations of these last two places are 1500 and 1538 m, respectively (Medem, 1965). Of 24 Colombian specimens with elevational data, all were from 200 m or below. Bothrops nasuta is both terrestrial and nocturnal, although specimens can be found sunning themselves by day (Smith, 1943; pers. observ.). The species can also be found on low bushes or shrubs (E. H. Taylor, 1954; Medem, 1968). Posada-Arango (1889b) mis- takenly believed that B. nasuta is ar- boreal, as did Amaral (1927b), when he said that B. brachystoma ( = B. nasuta) "is a tree viper. "Specimens have been found inside cavities in the trunks of trees, underneath roots of trees or rocks, on top of piles of rubbish (Alvarez del Toro, 1973), under rotting logs (Stuart, 1958), inside huts (Stuart, 1948), among loose rocks of ruins, (Smith, 1943), or simply on the forest floor (Duellman, 1963). We have collected B. nasuta un- der and lying near logs in the Turrialba area of Costa Rica. Picado (1931a) reported that new- bom Bothrops nasuta will eat earth- worms and later will consume small lizards of the genus Anolis. Amaral (1927b) stated that the species feeds on lizards and Alvarez del Toro (1973) indi- cated that it feeds on anoles and mice. Picado (1931a) reported an instance of cannibalism involving juveniles. We have observed cannibalism among adults and juveniles in captivity. We have also observed them to eat leopard frogs (Rana sp.), anoles (Anolis sp.), and mice in captivity. Bothrops nasuta has been reported to have 8 to 18 young (Picado, 1931a). A specimen in our collection (now LSUMZ 36898) from near Turrialba, Costa Rica, gave birth to 14 young in September, 1967. Bites from B. nasuta are known to cause fatalities in human beings (Daniel, 1949), although Minton and Minton ( 1969) did not believe they are capable of doing so. ANALYSIS OF VARIATION Twenty-six characters were analyzed for geographic, sexual, ontogenetic, and individual variation. These characters are discussed below. Intemasals. - Typically, Bothrops 92 Tulane Studies in Zoology and Botany Vol. 22 nasuta has the intemasal scales in medial contact or separated by only one scale. One specimen (FMNH 68056), how- ever, has the intemasals separated by two scales. This specimen also has one shortened and one elongate intemasal scale. Two specimens (LACM 45413, 45416) from Colombia do not have the intemasal scales as elevated as do other B. nasuta, but are typical in all other respects. The frequency of medial contact of the intemasals varies geographically. All of the specimens of B. nasuta from Mexico southward to Nicaragua have their intemasals in mutual contact. Three specimens from Costa Rica and one from adjacent northwestem Panama have their intemasal scales separated by one scale, but two of these specimens are from the Peninsula de Osa. The fre- quency of the mutual intemasal contact is reduced in specimens from central Panama southward to Ecuador (Table 1). In addition, specimens from central Panama and Ecuador have the intemas- als in contact less frequently than do specimens from Colombia. In all of the specimens but one, which is a juvenile, from the Peninsula de Osa, the intema- sals are less elevated and the snout is more attenuate. Nasofrontals. - The numbers of nasofrontal scales in male B. nasuta av- erage slightly lower than in females (Table 2) but the differences are not statistically significant. Specimens from central Panam^ ex- hibit a high average number of nasofron- tal scales relative to other areas of the range (47.21 + 2.63 as opposed to 37.7910.89). Intercanthals. - Males average fewer intercanthals than do females (Table 3), except in central Panamd, but the latter differences will probably be reversed with study of additional material. Central and westem Panamanian specimens have a higher average number of intercanthals than do specimens from the remainder of the range (5.48±0.36 vs. 4.92±0.10). Interoculars. - The average number of interoculars is higher in females than in males in the entire sample but the differ- ences are not statistically significant (Table 3). However, specimens from the northem portion of the range (westem Panama northward), excepting the Peninsula de Osa, show no statistically significant average difference between the sexes, whereas those from that point southward do (Table 3). Interictals. - Generally, males have fewer interictals than do females but the differences are not statistically signific- ant (Table 3). Rostral. - Two specimens (AMNH 10981, UMMZ 91078) have the rostral scale horizontally divided. All other specimens have a single rostral scale. Nasorostral. - A nasorostral scale is occasionally present in specimens from disparate parts of the range. The inci- dence of females possessing this scale appears to be greater than that of the males, inasmuch as ten of the twelve specimens having 1 or 2 nasorostrals are females. Nasal. - All specimens of B. nasuta examined have a partially divided nasal scale, except for two, which have the scale completely divided. The usual con- dition is that of a partial crease at the upper and lower sections of the scale but a few specimens exhibit a complete divi- sion of this scale above or below the naris. Prefoveals. - One specimen (UMMZ 117735) has the prefoveal scales extend- ing to the rostral scale thus excluding the nasal scale from contact with the first supralabial. Females have more prefoveals on the average than do males throughout the range (Table 4). Subfoveals. - With respect to the sam- ple as a whole, males have significantly fewer subfoveals than do females (Table 4). Preoculars. - There are generally an upper, middle, and a lower preocular No. 2 Hognose Viper 93 Table 1. Variation in internasals, body blotches, and sex ratios in Bothrops nasuta (parenthetical numbers in first and last columns indicate sample size; mid- dle columns indicate range, mean, confidence limits, and sample size, where applicable) Internasals Body blotches Males Females Sex ratios % in contact % of females Mexico 100(2) - 17 100(2) Guatemala 100(13) 18-22(19.29+1.16)7 15-17(15.75+0.74)8 61.1(18) Honduras 100(16) 17-20(18.50+19.06)2 15-18(15.88+0.94)8 81.3(16) Nicaragua 100(7) 17-19(17.67+2.88)3 16 57.1(7) Costa Rica 98.9(95) 16-22(18.36+0.63)25 15-20(17.22+0.48)36 60.0(95) Western Panama' 90.0(10) 19-22(20.00+4.30)3 16-19(18.20+1.04)5 70.0(10) (Northern Population) 98.6(148) 16-22(18.60+0.49)40 15-20(16.90+0.33)60 63.5(148) (Osa Population) 71.4(7) 18-20(19.0+12.71)2 15-19(16.67+5.18)3 57.1(7) Central Panama' 22.2(18) 19-23(19.00+4.68)4 16-19(17.77+0.70)13 78.9(19) Colombia 66.7(57) 15-23(19.31+0.77)26 15-22(18.61+0.66)25 52.6(57) Ecuador 22.2(9) 17-20(19.00+1.52)5 17-18(17.75+0.80)4 44.4(9) (Southern Population) 52.4(84) 15-23(19.23+0.66)35 15-22(18.26+0.45)42 57.6(85) Totals 81.6(239) 15-23(18.90+0.39)77 15-22(17.44+0.29)105 61.3(240) (95.0% of the specimens examined). The contact of the preoculars with the orbit is inconsistent, expecially since there is often a fleshy rim of tf«ue in- terposed between the preoculars and the orbit. Individual specimens exhibit differ- ent preocular arrangements on either side of the head. There are occasional specimens that do not have a middle preocular. This condition usually occurs when the upper and lower preoculars are enlarged and in contact with each other, thus excluding the middle one from orbi- tal contact. In one sp)ecimen (MCZ 55069) an enlarged supralacunal takes the place of the middle preocular. Soboculan and Postoculars. - The subocular and postocular scales vary in number (see description). In many speci- mens counts varied widely from one side of the head to the other. Specimens had as few suboculars as one or as many as four and postoculars range from two to four on either side of the head of the same animal. Ocnlabiab. - The usual number of oculabials present in B. nasuta is two (88.8% of specimens examined). Male B. nasuta from the entire range have significantly fewer oculabial scales than do females (Table 5). The number of specimens in which the 94 Tulane Studies in Zoology and Botany Vol. 22 c u ■o to s- E 3 n3 0) S- 3 l/l (O c l/l Q- O i- x: +-> o CO (D ,— Xi 0 c •r— 0 r— s- a. 4- Q. 0 ro 01 ro (U C s- OJ 1/1 M ^— »t— to m -<3 =J O) ro f^ 0 Q- U3 3 i to to M ■0 to C -*-> OJ •r- > E •f— c r^ •n- q; C 0 0 c (U 4J T3 <0 • r— 4- ^ c (O 0 > u CvJ QJ !q CM CO «■ ^. ir> in CM LO U3 <^. r^. <^, ^ <» + + 1 + 1 + 1 + 00 CM 0 lf> VO C71 liO 0 CO r^ ^ CO LD 0 ro m ro n ^3- ro CO 10 ^ \o ^3- «T ^a- m ^ o CO Si *?! + 1 o CM f— f— IT) VO 0 ^r ro r^ 00 U3. 0. 0. p_ + + + 1 + 1 0 ^a- 0 U1 lO CJ^ 0 CJ^ en r^ co' CM CM CM CM 0 CM 0 cn CO CO ro CM ■— . o. ■— , U3 CO CSJ VO CO CO ro p^ CO r^. M 00 00 • r^ «r • f^. ^, 0, CM A' &' ^' 4 ro iCl VD 0 •— r- «»• Ti Ti 5;. r^ r~ CD CO ro «* lO O, ro, in Qt CO CM CO (M CM <7> CT> I ±1 0 CO CO 0 VO ro CO ?1 ro VO 0 d •a- ro ro Cv) C •0 a K c (- lO u QJ 3 i. <0 Ol ■*-» T3 to 4J 4-» 10 C u bO m 3 0 •^■ 0 HI tJ3 :c z <_> 3 I' to d, *' CO CO ir> CO CM I' e*l CM Q. O a. c 1. 01 o z ♦ I in I' o «3- + \ VO VO c o 0 «r ro CO VO in 0 '^i + 1 + 1 Vl VO C7^ ?l in CM o m CO CO m •a- CO CM 0 VO VO CJ> i 0 i 0 »a- CO f-^ CM CO cn CM TT vn ro 0 ,_; •^i A' i' 00 LO ?l CO CM ?1 ±1 to ro O CM 1' CO CO CM •a- CM CO ?l r^ CM CM ro 4' o r— »— r^ vn CO o CM "a- CO in CSJ CM cn I' CO C7^ CM CO CM .— ■•- 1- c o Q. O O. o in e o en 0) JQ CJ ^- cr> r* «3- *— * ^— ^— «a- in r^ CT> r^ o ro in r. vo a> • r^ r^ o »»■ CM CSJ • • • ^— o, o, '^i cj, ^~ d, + 1 + + S' -t-l -t-l + o C71 in en co' o o CO m in CM i£> ID lO «j- in >a- «^ •a- CM CM CM CM CM CM CM r-. 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JZ +J o CD c n] o c > o > o c o I. J3 o o o, o, +1 +1 +1 +1 o C3 r^ ^ o o ^ •— ?l o o E CM 1 — . — «» Lf> r^ o CO CM CM CO ^ in U3 5' «3- en CM O) > o CM o o CO ?1 o o, o, + -t-l o% n o CO <— CM >— .— <0 E P— CO «3- ir> r-~ CM *— ^ ^^ - — . CM CO O to lO o o\ CM O CM «? LT) > . r— CD, , — '^l o, o, + 1 + 1 + + + + ro «r o ro o ^— , (£) o r^ o . . . > 00 VD to IT) IT) lO — * >^_^ — ' w— CM o >£l 00 cr> •— •— CO «» «T Lr> n ^ 01 s- Q. o, ^, r^ o. o + + + 1 + + «»■ o 1-^ CM r^ •— 00 •— VO >— n »»• VO '3- vo 1- 00 00 cn r-~ o u >— l/l D -.- "0 m C71 cr E I- -o I CM c o 3 Cl O o. c I. Ol o en ro ?1 in o ?1 in O ^ CO ^ CJl CM in • — •~ •— C3, o, o + + + ro o in ^ ro CM cn o o. in «3- 00 o ?l CM lO «3- CM cn C3 + 1 o + 1 o o in C7^ ?l c o 3 a. o c^ ?1 to CO .— CM .— T CM in «a- CO 00 m •— ^r ci, o, d + + 1 + CO cn o ^ c o CM CM CM CO CO ro CM CM in in ro CM CO o in o 1 — CO f — . ^ — . •q- CO in r^ «j- cn CO d, d, + + 1 d. CO r^ + CM CM o in in ID • cn 'J lO ro O .— .— CO rn CM CO in ^' lO C7^ CTi ID «3- CO CO <7l m i£> ro lO CO ?l en o CO CM CT> 00 O ?1 lO lO CM CM C7< CM CM 1- I CM cr> »r ro •>* ^ 1 — ( — "3- ro CO r^ m ro en »!■ esi »^ »— ^ o r^ o 'd- o ■<3- o O o, o, o, o, o. O, o. o. o. o. o. o. d, + 1 + 1 + 1 + + 1 + + 1 + 1 + + -t- + 1 -4- o o CM o «T o CM in r^ o CO •^ lO o o cn o o o O CM CJl CSJ CO o Csl CM ^ CM CM CM CM CM ^ CJ CM CM CM CM CM CM CM CO CM m CO CM nr CO "a- «3- CM CO CM ^ r^ CM LD CM CO m CO LD «» eg n en CO r^ CO r- Ln fO , — LO r^ U3 CM o CO o ro "a- r— in 1 — o o, CD, O, CD, o. o. O. o. o. o. o, o, + + + 1 + 1 + + + + + 1 -t- + 1 + o O ro C^ o o o o ^ o LD CM 1 o ^O 00 CO o cr> o o en o en o\ CM , — , — ,J CM ^ CM CM r— CM ^ ^ CM CM CM m CM CO CM ro CO CM ro CO "■ "■ '' '^ '' r^ w— n r^ ■tj- in o en «J- CM f— •— ^ LT) r^ cn «r *— ro ^ ■<3- »— O 00 00 cr> CM O O in ^ ro" ro ro 00 00 tNl CM ro •— in •— •^ CM »— *3- ' — o o O. O. CD, CD, CD, CD, o, O. CD, CD, d. d. + 1 + 1 + 1 + 1 + 1 + 1 + 1 + + + 1 + + + IT) r^ CM LO >£. lO Ln o O ro ro CO ro CM r^ tj\ CM r^ CO r-- o LO r-^ LO LD rN. CVJ f— , CM ,_ o ' CM ^ ,-i ._■ ,Ji ,-1 f— , — f — 1 — I — , — f~ r— 1 — 1 — f— 1 — r— O ro ro PO ro CM ro ro CM ro CM ro CO r- »— •— »— *— •— < — »— '— '— *— • — 1 CSl 1 t o CM O 00 o O ^ O o ^ CD o r^ m ro ro ro Ln ro ■3- CM LO ro in ro en n *r r^ ^o •a- -3- LO cn LD . — , . — . ^— ^ m o^ ID LO • CO CO *~ "J- r^ ^~ c o C. CD. CD, o, o. c, • + 1 + 1 + + 1 + 1 + 1 + 1 + + 1 CD, o o, o o ^3- CO Ln r^ 00 Ln + + + 1 , CM o LO O ro o LO ro LD O ro CO LO en 00 r— , — .— 1 — O . — CD , — CD • •— <— »— »— ■ — »— '— •~ d o o CM CM CM CM " CM CM ro CM d. 3 CI. ■— "— ■— 1 '~ I '~ I r ^_ ro ro o o . — c o O o O O 1 1 1 •" •" *" '~ *"* ' ' '~~ CTi en en CM r^ ,^ m r^. «3- ^ Ln o en «» O , <3- m r^ Ol r— ro ^ «* r— CO Ln CM o CT^ .— , — CT. '^ 1 — ro (T> o r^ ?, CM ro m o ro O ?1 CM ■-; ro """_ o + 1 in O O O, CD, d, O, CD, CD d * CM Ln o o 'o- LO in in CM in LD o • • v_^ oi CT^ o^ 0^ o\ <^ »— 0^ as en en en ,_ O , — o r~ o 1 — r— O o o o r— 1 >— -— »— »— '-' •~ •~ •~ •"■ o CT^ 1 CJ^ c^ CO CD CTi 00 CT. CO en en CO LO 00 CO r^ CO ro CO Ln ro <* CM Ln ro en uo «T O ro CD LD o CM ro O LO CM in fj ^ O LO ro •-; CM — ; o O d O o CD, CD, CD, CD, d d, d, -t-l + \ o + 1 + 1 + 1 + 1 + 1 + 1 + 1 + 1 i^ m' -M CM r^ O o ro in o ^ 00 1 CM CO o CM r— CM o •— *— '~ ^~ '^ cy^ CT> o O^ Ol m en c^ CT> en en 0^ (7> 1 E lO c 1 C o Q. O Ci. 1 00 c o ♦J 1 en c 10 1 00 1 HI 4-» 1- ra 1. ■4-J § -C3 •0 3 o X (Q c u (/> (/) o (/I c 3 o <1J 3 o o 41 z o Oi o U w^ 3E o I z CJ Zf (_) <_) LlJ V— 98 Tulane Studies in Zoology and Botany Vol. 22 oculabials are more than two is greater in Colombian and Ecuadorian specimens (21.2%) than in the remainder of the range (4.1%). Supralabials and Infralabials. - The numbers of supralabials and infralabials average more in females than in males over the entire range (Table 5). All but one specimen (UMMZ 91078) have the first infralabials in contact. Ventrals. - The number of ventral scales is slightly, but significantly higher in females than in males (Table 2) con- sidering the range as a whole. A rela- tively high number of ventral scales is present in specimens of B. nasuta from western Panama northward to Mexico (129-145, x = 137.68 ±0.42). A relatively low number exists in specimens from central Panama southward to Ecuador (123-143, 7= 132. 15 ±2.58). Subcaudals. - As is usual in snakes, the subcaudals are more numerous in males than in females (Table 2), cor- relating with their greater relative tail length (Table 7). Dorsal Scaie Rows. - The dorsal scale rows usually number 25-23-19 (see de- scription). Specimens from the Pem'n- sula de Osa are unusual, however, in possessing atypically high dorsal scale row counts (Table 6). Only one of the seven specimens available from there has the above-mentioned count. Two specimens have 25-25-19 rows, three have 27-25-19 rows, and a sixth has 29- 27-19 rows. The latter specimen has the highest neck and midbody counts of any specimen examined. Canthals. - All of the specimens that possess more than one canthal are from central Panama. Of 19 specimens from this area, 7 have two canthals on at least one side of the head; three specimens have two on both sides of the head. Postcanthals. - Females show a higher incidence of two postcanthals (instead of one) than do males throughout the range (Table 4), with the exception of the sample from central Panama. Viewed over the entire range, the differ- ences are significant. Color and Pattern. - Various aspects of color and pattern in B. nasuta vary individually, ontogenetically, sexually, and geographically. There is considerable intrapopula- tional variation with some individuals being brightly marked on the head and body, whereas others are almost uni- color (see description). The only series of specimens from a single locality from which adequate notes on ontogenetic color variation could be taken consists of twenty speci- mens from the Rio Patia in southern Co- lombia (AMNH 107912-13, 109794-811). The juvenile specimens are noticeably brighter in color and more distinct in pattern than are the adults. In larger specimens the brownish gray ground color is basically the same but the brownish black to black dorsal blotches are faded and the whitish border of these blotches is indistinct. The fading of the pattern begins on the lateral portion of the dorsum and proceeds dorsad. The vertebral pale line is bright in the young and gradually fades in larger specimens, particularly females, until, in some specimens the line disappears altogeth- er. In juveniles the dorsal blotches are discrete and well-defined, except for the first two or three blotches, which are reduced to small spots. As the animals increase in size, the blotches gradually fragment into two portions, which are laterally displaced from the middorsal line. This fragmentation and lateral disp- lacement of the dorsal blotches proceeds from anterior to posterior. Males generally possess a more con- trasting facial and dorsal coloration than do females, although occasional dark male specimens are also seen. Males av- erage more body blotches than do fe- males and the values for the entire sam- ple are significantly different (Table 1). A living specimen from Ecuador in the senior author's private collection ex- hibits a peculiar temporary coloration prior to ecdysis. The animal is grayish No. 2 Hognose Viper 99 Table 6. Variation in dorsal scale rows in Bothrops nasuta (explanations as for Table 2). Males Females Mexico Guatemala Honduras Nicaragua Costa Rica Western Panama 25(25.00+0)7 23(23.00+0)7 19-20(19.15+0.35)7 25(25.00+0)2 23(23.00+0)2 19(19.00+0)2 25(25.00+0)3 23 23.00+0)3 19(19.00+0)3 23-27(24.92+0.52)39 21-23(22.81+0.20)39 19-20(19.03+0.32)39 25(25.00+0)3 23(23.00+0)3 19(19.00+0)3 25(25.00+0)2 23(23.00+0)2 19(19.00+0)2 25-27(25.19+0.41)11 23(23.00+0)11 19(19,00+0)11 25-27(25.36+0.49)12 23-25(23,18+0.37)12 19(19.00+0)12 25(25.00+0)4 23(23.00+0)4 19(19.00+0)4 23-27(25.08+0.14)57 21-25(22.99+0.11)57 17-21(19.01+0.10)57 23-25(24.69+0.86)6 23(23.00+0)7 19(19.00+0)7 (Northern Population) 23-27(24.93+0.18)54 21-23(22.85+0.14)54 19-20(19,04+0,05)54 23-27(25,09+0,12)92 21-25(23.02+0,08)93 17-21(19.00+0,06)93 (Osa Population) 25-27(25.70+2.87)3 23-25(24.37+2.87)3 19(19.00+0)3 25-29(27.07+2.60)4 25-27(25.53+1.60)4 19(19.00+0)4 Central Panama Colombia Ecuador 23-25(24.53+1.60)4 22-23(22.68+1.43)3 19(19,00+0)4 25(25.00+0)27 23(23.00+0)27 19(19.00+0)27 23-25(24.63+1.11)5 21-23(22.63+1.11)5 19(19.00+0)5 23-25(24.75+0.39)15 23(23.00+0)15 19(19.00+0)15 23-27(25.44+0,36)30 23-25(23,22+0,11)30 19-21(19.12+0.56)30 25-27(25.53+1.59)4 23(23.00+0)4 19-20(19.26+0.80)4 (Southern Population) 23-25(24.89+0.02)36 21-23(22.91+0.13)35 19(19.00+0)36 23-27(25.20+0.27)49 23-25 23.13+0.14)49 19-21(19.09+0.13)49 Totals 23-27(24.94+0.13)93 21-25(22.92+0.12 92 19-20 19.02+0.13)93 23-29(25.18+0.13)145 21-27 23.12+0.10)146 17-21(19.03+0.06)146 100 Tulane Studies in Zoology and Botany Vol. 22 brown dorsally and powder gray ventrally. During the premolt period the ventrals and the first dorsal scale row and the ventral half of the second row turn a bright silver, contrasting markedly with the color of the remainder of the dorsum (Fig. 3). Size and Proportions. - The average dimensions in head length, head width, head height, total length, and body length are greater in females than in males (Table 7; Fig. 4). Males possess a greater average relative tail length (Table 7), correlated with the higher av- erage number of subcaudals (Table 2). Female specimens appear to be stouter than do males, but this feature is difficult to quantify because of variation caused by preservation. Sex Ratios. - We sexed 240 specimens of Bothrops nasuta. A prevalence of females is present in collections from Mexico southward through central Panama (64.9% females vs. 35.1% males). The sex ratios in samples from Colombia and Ecuador are more evenly balanced (51.5% females vs. 48.5% males) (Table 1). Considering the number of specimens involved, it seems unlikely that these differences result sim- ply from artifacts of collecting. The reasons for the greater percentage of females in samples from Panama north- ward, however, remain obscure. ^ vi^N^'s/^^-N^' Figure 3. Section of body of a female Bothrops nasuta from Ecuador illustrating the marked con- trast between the lower ventral color and that of the dorsum that is exhibited prior to ecdysis. Figure 4. Adult male (lower) and female (upper) Bothrops nasuta from northern Honduras illustrat- ing sexual dimorphism in size and the typical color pattern of members of the northern population. DISCUSSION On the basis of the foregoing analysis, the following characteristics have been shown to exhibit geographic variation in Bothrops nasuta: intemasals; nasofron- tal; intercanthals; interoculars; ocu- labials; ventrals; dorsal scale rows; can- thals; color and pattern; sex ratios. These characteristics show differing patterns of variation. The frequency of mutual intemasal contact decreases markedly between samples from western and central Panama. Ventral numbers are highest in the northern section of the range (western Panamd northward) and lowest in the southern section (central Panama southward). Average numbers of interoculars are not significantly diffe- rent between the sexes in samples from western Panama northward, whereas the differenes are significant from that point southward. Central Panamanian speci- mens are distinctive in having a higher average number of nasofrontal, as well as being the only specimens jxjssessing more than one canthal scale. Panaman- ian specimens have a higher average number of intercanthals than do speci- mens from the rest of the range. The preponderance of female specimens from the northern portion of the range south to and including central Panama is No. 2 Hognose Viper 101 Table 7. Sexually dimorphic characters of size and proportions in Bothrops nasuta (measurements in millimeters - parenthetical numbers indicate range, mean, and sample size). Males Head length Head width Head height Total length Tail length Body length Tail/total length ratio 11-24(17.0)93 7-15(11.3)93 4-10(6.6)93 162-463(291.1)88 20-62(36.2)88 142-401(254.9)92 0.103-0.137(0.124)88 Females 12-37(22.6)147 8-28(14.9)147 4.14(8.6)147 156-635(349.0)139 16-64(35.7)139 140-571(313.4)142 0.092-0.122(0.103)139 in marked contrast to the more even sex ratio in samples from Colombia and Ecuador. Specimens from Colombia and Ecuador have higher average numbers of oculabials than do specimens from the remainder of the range. The Peninsula de Osa specimens are distinctive due to: (1) a higher number of dorsal scale rows on the neck and at midbody; (2) the juvenile tail color persisting into adult- hood; (3) the shape and degree of eleva- tion of the intemasal scales, and (4) a distinctive dorsal body coloration. Obviously, the most distinctive speci- mens are those occurring in the Penin- sula de Osa. Due to the small number of specimens available from this area, how- ever, it remains an area in need of critical study. Other, less distinctive geographi- cal groupings may be circumscribed as well. The patterns of variation in the samples of populations of B. nasuta out- side of the Peninsula de Osa are such that no clear cut distinctions can be drawn (i.e., the patterns of variation are dis- cordant). On the other hand, the pattern of variation in several characters (frequency of mutual intemasal contact; number of ventrals; nasofrontal, can- thals, and intercanthals; sexual differences in average numbers of interoculars; and sex ratios) is such that a number of changes in those patterns occur in Panamd, with the most obvious changes occuring between western and central Panamd. Specimens from central Panama, therefore, in general resemble specimens from South America, whereas those from western Panama' resemble specimens to the north (excluding those from the Osa Peninsula). On the above basis, therefore, we pre- fer to recognize three distinctive popula- tions of the hognose viper, Bothrops nasuta: the northern, Osa, and southern populations. Inasmuch as these popula- tions do not exhibit equivalent levels of divergence, we do not recognize infra- specific taxa. We feel that to do so would tend to obscure the complex relation- ships we have detailed above. The popu- lations are characterized below. The Northern Population. - This population extends from southern Mex- ico southward to the Boca de Almirante area of western Panama, with the excep- tion of the Peninsula de Osa-Golfito area of southern Costa Rica. This population is characterized by having a greater in- cidence in the mutual contact of the inter- nasal scales (98.6% of the specimens ex- amined), a higher average number of ventral scales (males, 129-143, x = 136.98; females, 128-145, x=138.14), a lower average number of nasofrontal scales (males, 22-46, x*= 33.77; females, 25-54, x=35.78), and a lower mean number of oculabials (%.6% of specimens having two or fewer). Typical examples from this population are iUustrated in Figure 4. The Osa Population. - This population is confined to the Peninsula de Osa-Golfito area of southern Costa Rica and possibly adjacent western Panama. This population is characterized by having a higher mean number of dorsal scale rows at the neck and midbody, the tendency of juvenile tail color to persist into adulthood (especially on the under- side of the tail), a lesser degree of eleva- 102 Tulane Studies in Zoology and Botany Vol. 22 tion of the anterior portion of the in- temasal scales, and a tendency for pos- session of transverse body bands (Fig. 5), as opposed to alternating blotches or spots. The Southern Population. - This population occurs from Valle de Anton and the Canal Zone in central Panama southward to Ecuador. This population is characterized by having a relatively low incidence of mutual contact of the internasal scales (52.4% of specimens examined), a lower average number of ventral scales (males, 123-141, x = 130.57; females, 127-143, x= 133.35), a higher average number of nasofrontal scales (males, 25-56, x=40.14; females, 29-59, x=42.84), and a higher mean number of oculabials (17.6% of specimens having three or more). Biogeographic History. - Recently several authors have shown that alter- nating dry and humid climatic periods and sea level fluctuations occurring dur- ing the Pleistocene have had a marked effect on the evolution and distribution of Neotropical vertebrates (van der Hammen, 1961; Haffer, 1969; Vuilleu- mier, 1971; Duellman, 1979). In addi- tion, several recent taxonomic studies on Bl 1 ^^^^^^1^ * W ^m^^ Figure 5. Adult female Rothrops mmaa (USC- CRE 6391) from Rincon de Osa. Puntarenas Pro- vince, Costa Rica, illustrating the dorsal pattern of transverse bands. South American and Central American amphibians and reptiles have shown a close correlation between evolutionary events in the group in question and the paleogeographic events detailed in the above-cited papers (Vanzolini and Wil- liams, 70; Duellmn, 1972; Heyer, 1973; Duellmn and Crump, 1974; Wilson and Mena 1980). Although the evolutionary relation- ships of the Bothrops lansbergi group, to which B. nasuta belongs, are imperfectly known. Burger (1971) expressed the opin- ion that the members of the lansbergi group, as well as those of the godmani and nummifera groups constitute a dis- tinct genus, Porthidium. He stated that "the lansbergi and nummifera groups di- verged independently from the godmani group." In addition, he opined that ^^ Porthidium nasutum, which is similar to P. godmani in some respects and specialized in others, is close to the ancestry of the lansbergi group." For these reasons we assume, for the present, that the point of origin of the species B. nasuta lies somewhere in Nuclear Cen- tral America and, most likely, southern Mexico. On the basis of information in the above-cited papers, we postulate the fol- lowing sequence of events to account for the evolutionary history of Bothrops nasuta: 1. Movement of the B. nasuta stock from the north (probably southern Mexico) into Central America during the upper Pliocene along the eastern humid corridor. 2. Movement of the stock of the Osa population into southern Pacific Costa Rica through low plateaus and/ or narrow river canyons in the uplift- ing Talamanca range during the turn of the Pliocene-Pleistocene. 3. Isolation of the Osa population in the Peninsula de Osa by rising sea levels during the interglacial periods of the Pleistocene. No. Hognose Viper 103 4. Isolation of the northern and south- ern populations in wet forest refugia in Caribbean Central America and the Pacific side of Colombia, respect- ively, during a period of dry forest expansion in the Quaternary. SPECIMENS EXAMINED Locality data for each specimen ex- amined are listed below. The data are arranged as follows: alphabetically by country, state (department or province), and locality; alphabetically by the first letter in the abbreviations for the mu- seums, and numerically after each mu- seum abbreviation. Specimens lacking precise locality data are listed in the first most restricted political unit possible. Localities enclosed by quotation marks are not mapped. The localities and specimens are: COLOMBIA: "Boca de la Raspadura," AMNH 18298-300. Anti- oquia: "Medellin," AMNH 35735; Rio Arquia, Belen, LACM 45413; Sabanalarga, Cauca Valley, AMNH 35795. Cauca: Quebrada Guanguf, 0.5 km above Rio Patia, 100-200 m, AMNH 109794-811; Quebrada Guangui, Rio Patia, 100 m, AMNH 107912-13. Choco: "no specific locality," AMNH 8067-68; Andagoya, MCZ 29255, USNM 124259, Andagoya-Condota area, UMNZ 121043-44; be- tween Andagoya and Condota, UMMZ 121058; vie. Playa de Oro, upper Rio San Juan, 200 m, AMNH 108460; Quebrada Taparal, 20 km N Falestina, CAS 119919; Ri'o Atrato, S of Quibdo, USNM 140050; Ri'o San Juan, USNM, 72352; Sierra de Baudo, ANSP 25573-78; trail between Quebrada Bochorama and Ri'o Tadocito, LACM 45416. Valle: Buenaventura, at Rockerfeller Lab, 12 m, TU 18712; km 13 from Buenaventura to Rio Calima, 45 m, FMNH 165478, 165893; near Buenaventura on land of Carton de Colon, TU 1871 1; Rio Calima, Quebrada de la Brea, USNM 124260; Rio Calima, 7 km from lumber camp, FMNH 165492, 165495, 165566, 165725-26, 165900; Rio Paposo, Virology Field Station, USNM 151711-12. "COLOMBIA or ECUADOR": USNM 22422. COSTA RICA: "Hospital San Juan de Dios," KU 34636; "Costa Rica,"UF 20627, 30709, UMMZ 133893, 133911. Alajuela. Canton de San Carlos, Muelle San Carlos, 65 m, USC-CRE 271 1 ; Cariblanco, UCR 1431; Cataratas de San Ramon, Lado San Carlos, USC-CRE 2754; Ciudad Quesada, 656 m, USC-CRE 2693; Ciudad Quesada, San Roque, 580 m, USC-CRE 2695; San Carlos, FMNH 191789, UMMZ 13 1327-29; Sarapi- qui, 100m, UCR 109, 2990, USC-CRE 2691. Cartago: Pavones, nr. Turrialba, 819 m, KU 140087, LSUMZ 36898-99, USC-CRE 2710; Tur- rialba, 624 m, KU 30982-87, 34876, 34879-80 35734-35, MCZ 55067-74, UCR 1870; Turrialba, IICA, 600-624 m, AMNH 69722, FMNH 179083 KU 25689, 34635, 34637-38, MCZ 56116, USC- CRE 646; Turrialba, 3 km from IICA on road to Pavones, 630 m, USC-CRE 190. Guanacaste: El Silencio de Tilaran, 825-850 m, USC-CRE 6217; 5 km ENE Tilaran, 600 m, KU 36693; Tilaran, 560- 562 m, KU 35737-38, USC-CRE 2694, 2712, 7131, 7163. Heredia: "no specific locality," UF 30492, 30495, 30498, 31795-96; La Selva, central trail. 60 m, USC-CRE 8291. Limon: La Lola, 39 m, KU 34005, UMMZ 117736-37, USC-CRE 127 (2 spec), 128 (3 spec), 140 (2 spec), 162, 203, 207, 655 (2 spec); Limon, MCZ 19744; Los Diamantes, UMMZ 117738; nr. Penshurst, HSH/RSS 600; Penshurst, UCR 2721-22, 2738-39, 2940-41 ; Puerto Viejo, UCR 162; 4 km E Puerto Viejo, UCR 308; Suretka, nr. Mt. Mirador, KU 35736; Sipurio, Talamanca, AMNH 17332; Zent, MCZ 11457-58. Puntarenas: "no specific locality." UMMZ 83185; Golfito, 5 m, USC-CRE 836; 6.7 km E Golfito,' LSUMZ 1 1652; RincondeOsa, LACM 114153-54; vicinity of Rincon de Osa, 20-40 m, USC-CRE 6391; Rincon de Osa, Camino del Pacifico, UCR 3310, 3359. San Jose: Puriscal, UCR 110. ECUADOR: Esmeraldas: Playa de Oro, Santi- ago River, USNM 20625-28; Quininde', USNM 165317; Ri'o Capapas, MCZ 11169. Pichincha: 18 km W Santo Domingo de los Colorados on Chono road, USNM 165320; 5 km W Santo Domingos de los Colorados, USNM 165321; 12 km NW Santo Domingo de los Colorados on Quininde' Road, USNM 165319. i^inm loojiy. GUATEMALA: Alta Verapaz: Finca Chama, UMMZ 91077-78. El Peten: 12 km NW Chinaja, 130 m, KU 55705; PiedrasNegras, USNM 110415; Sayaxche', UCM 22367; Sojio (=Toocog), AMNH 69966-67, 69987; Tikal, 283 m, MCZ 55436-37, UF 13866, 13868-69, UMMZ 117944; 4.8 km S Tikal, KU 157665; 13.5 mi S Tikal, KU 157664; Valon- tun, 4 km SE Tikal, AMNH 100410. Izabal: 12.8 km SE Cayo Piedra, 153 m, ANSP 22149. HONDURAS: Atldntida: Dakota, East Line, MCZ 20247; Jilamo, MCZ 34385; Lancetilla, AMNH 46958-60, MCZ 38781 ; Sonora Farm, Tau- jica District, MCZ 20493; Tela, AMNH 46961; Tela, Juliana Farm, MCZ 22023. Cort/s: La Cumbre, nr. San Pedro Sula, MCZ 32028; San Pedro Sula, MCZ 33332-33. Snata Barbara: Santa Barbara, MCZ 27566; Trinidad. MCZ 27565. Yoro: Progreso, MCZ 22024, 268722UMMZ 62522^ MEXICO: Chiapas: Lake Miramar, USNM 136966-67. "Veracruz," ANSP 4873. NICARAGUA: "vicinity of Poderoso," AMNH 12706. Matagalpa: "no specific locality," UMMZ 57654; Hacienda La Cumplida, 104 Tulane Studies in Zoology and Botany Vol. 22 900m, UMMZ 117735; 19 km N Matagalpa, UMMZ 116523. Rio San Juan: Los Sabalos, San Juan River, AMNH 28355. Zelaya: Eden Mine, AMNH 7411; Rio Huahuashan, Papel Camp, AMNH 70247. PANAMA: "Panama City," MCZ 37115-16. Bocasdel Toro: Almirante, FMNH 83466, 153847, KU 80247, 112597; 11 km NW Almirante, FMNH 153849, 153851-52, 153999; 12 km NW Almirante, UU 5564. Canal Zone: Chico Limpio Divide, 470 m, MCZ 42767; Madden Dam, ANSP 23889; nr. Miramar River, 183 m, MCZ 42768; Pequeni- Esperanza Ridge, 214-610 m, MCZ 42772-81. Co- de: El Valle de Anton, 550 m, KU 1 12596; El Valle de Anton, Finca Acre, FMNH 68054-56. Panama: nr. Boqueron, Candelaria, and Peluca Station, AMNH 68850. LITERATURE RECORDS BELIZE: Xuanantunich (Neill, 1965). COLOMBIA: Antioquia: Segovia (Nic^foro Maria, 1942); "Urrao" (Niceforo Maria, 1938, 1942); Zea District (Posada-Arango, 1889). Caldas: "Pueblo Rico" (Niceforo Maria, 1938, 1942). Choco: Cabeceras (Rendahl and Vestergren, 1941); Quibdo (Niceforo Maria, 1942. GUATEMALA: Alta Verapaz: Panzos (Bocourt, 1868). MEXICO: Chiapas: Rancho Alejandria, Municipio de Juarez (Alvarez del Toro, 1973). NICARAGUA: "Hacienda de Jericho [=Jerico], 991 m" (Gunther, 1895; Boulenger 1896); "Chontales" (Gunther, 1895; Boulenger, 1896). Rio San Juan: Greytown (Amaral, 1929a). PANAMA: Canal Zone: "Cerro Bruja" (Amaral, 1929a; Schmidt, 1933; Dunn and Bailey, 1939). ACKNOWLEDGMENTS We would like to thank the following people for allowing us to examine mate- rial in their care: Walter Auffenberg, Florida State Museum, University of Florida (UF); William E. Duellman and Julian C. Lee, Museum of Natural History, University of Kansas (KU); Harold A. Dundee, Tulane University (TU); Herbert S. Harris and Robert S. Simmons, Natural History Society of Maryland (HSH/RSS); Arnold G. Kluge, University of Michigan, Museum of Zoology (UMMZ); John M. Legler -Stated to have been from Progreso, Prov. Chiri- qui, Panama by Loveridge (1928) and Amaral (1929a). and Robert M. Winolur, Museum of Zoology, University of Utah (UU); Alan E. Leviton, California Academy of Sciences (CAS); Hymen Marx, Field Museum of Natural History (FMNH); T. Paul Maslin, University of Colorado Museum (UCM); Edmond V. Malnate, Academy of Natural Sciences, Phila- delphia (ANSP); Douglas C. Robinson, Universidad de Costa Rica (UCR); Douglas A. Rossman, Museum of Zool- ogy, Louisiana State University (LSUMZ); Jay M. Savage, University of Southern California-Costa Rica Expedi- tion (USC-CRE); Ernest E. Williams, Museum of Comparative Zoology (MCZ); John W. Wright, Los Angeles County Museum of Natural History (LACM); George R. Zug, National Museum of Natural History (USNM); Richard G. Zweifel, American Museum of Natural History (AMNH). We would also like to thank Jay M. Savage for providing us with information concerning a locality in Costa Rica, Roy W. McDiarmid and Charles W. Myers for providing us with photographs of Bothrops nasuta, and Albert Schwartz for bringing to our attention the matter of the correct spelling of the specific name for the species. LITERATURE CITED ALVAREZ DEL TORO, M. 1973. Los reptiles de Chiapas. Segunda edicion. Tuxtla Gutier- rez, Chiapas, Mexico, Gobiemo del Estado. v + 178 pp. AMARAL, A. do. 1925. South American snakes in the collection of the United States National Museum. Proc. U. S. NaU. Mus. 67(24): 1-30. . 1927a. Studies of Neotropical Ophidia. V. Notes on Bothrops lansbfrgii and B. brachystoma. Bull. Antivenin Inst. Amer. 1(1): 22. 1927b. Studies of Neotropical Ophidia. VII. An interesting collection of snakes from west Colombia. Bull. Antivenin Inst. 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CORKUM Department of Zoology and Physiology , Louisiana State University, Baton Rouge, Louisiana 70803 ABSTRACT The following trematodes were collected from mammals in southern Louisiana: Alaria mustelae Bosma, 1931 from the raccoon, Procyon lotor (Linn.); Fibricola cratera (Barker and Noll, 1915) Dubois, 1932 from raccoons, opossums (Didelphis virginiana Kerr), and a mink, Mustela vison Schreber; F. lucida (La Rue and Bosma, 1927) Dubois and Rausch, 1950 from a raccoon and a gray fox, Urocyon cinereoargenteus (Schreber); Pharvngostomoides procyonis Harkema, 1942 from the raccoon; Heterohilharzia americana Price, 1929 from raccoons and opossums; Apo- phallus venustus (Ransom, 1920) Cameron, 1936 from raccoons; Parametorchis complexus (Stiles and Hassai, 1894) Skrjabin, 1913 from raccoons; Baschkirovitrema incrassatum (Dies., 1850) Skrja- bin, 1944 from an otter, Lutra canadensis (Schreber); Brachylaima virginianum Dickerson, 1930 from opossums; Gyrosoma singulare Byrd, Bogitsh, and Maples, 1961 from raccoons; Has- stilesia texensis Chandler, 1929 from swamp rab- bits, Sylvilagus aquaticus (Bachman); Maritre- minoides nettae (Gower, 1938) Rankin, 1939 from raccoons; and Rhopalias macracanthus Chandler, 1932 from opossums. Brachylaima virginianum, Fibricola cratera, F. lucida, Hasstilesia texensis, Heterohilharzia americana, and Rhopalias macra- canthus have been previously reported from Louisiana; the other species are new locality re- cords. The raccoon is a new host record for adult Alaria mustelae. Diagnoses are presented for species representing state records along with pertinent notes on the epidemiology and zoo- geography of each. INTRODUCTION During our studies on the life cycle of Alaria marcianae (La Rue, 1917) Wal- ton, 1949 in Louisiana we had the op- portunity to examine other mammals for trematodes. All mammals reported here- in were collected in southern Louisiana from an area bordered on the west by the Atchafalaya basin and on the east by Lake Maurepas. The mammals were col- lected by leg traps set in and around cypress-tupelo swamps. The following mammals were exam- ined for trematodes, one mink, Mustela vison Schreber; one gray fox, Urocyon cinereoargenteus (Schreber); one striped skunk. Mephitis mephitis (Schreber); two river otters, Lutra canadensis (Schreber); two swamp rabbits, Sylvila- gus aquaticus (Bachman); twelve opos- sums, Didelphis virginiana Kerr; and thirty raccoons, Procyon lotor (Linn.). Trematodes were fixed in steaming 10% formalin and stained in Semichon's aceto-carmine. All measurements are in micrometers unless otherwise stated; means are followed by the ranges in parentheses. Line drawings were pre- pared with the aid of a microprojector. Representative specimens for which diagnoses are given were deposited in the Manter Laboratory, University of Nebraska State Museum. The following is a list of trematodes we found from mammals that have been previously reported from Louisiana: From the mink we found Fibricola crat- era (Barker and Noll, 1915) Dubois, 1932 in the small intestine; both swamp rabbits harbored thousands of Hasstil- esia texensis Chandler, 1929 in the small intestine; in opossums F. cratera and Rhopalias macracanthus Chandler, 1932 occurred in the small intestine of all twelve, Brachylaima virginianum Dick- EDITORIAL COMMITTEE FOR THIS PAPER: DR. RICHARD D. LUMSDEN, Professor of Biology, Tulane University, New Orleans, Louisiana 70118 DR. F. SOGANDARES-BERNAL, Professor of Biology, Southern Methodist University, Dallas, Texas 75275 109 no Tulane Studies in Zoology and Botany Vol. 22 erson, 1930 in the small intestine of eight, and Heterobilharzia americana Price, 1929 in the mesenteric venules of three; from gray fox we have a single specimen of F. lucida (La Rue and Bosma, 1927) Dubois and Rausch, 1950 from the small intestine; no trematodes were found in the skunk; and in the ra- coons we collected F. cratera and F. lucida from the small intestine often and one respectively, and H. americana in the mesenteric venules of four. Lumsden and Zischke (1961) ade- quately described Fibricola cratera, F. lucida, Hasstilesia texensis, and Brachy- laima virginianum from Louisiana mam- mals. Malek et al. (1961) studied Heterobilharzia americana in southern Louisiana and found the raccoon to be the principal definitive host. Kaplan (1964) was the first to report H. americana from the opossum and com- mented on its rarity in marsupials. In addition to these previously reported trematodes we have identified seven species, one from the otters and six from the raccoons, that have not been re- ported from Louisiana and are of im- portance from either epidemiological or zoogeographical standpoints. Family DIPLOSTOMIDAE Poirier, 1886 Alaria mustelae Bosma, 1931 (Figure 1) Synonyms: Alaria freundi Sprehn, 1932; A. intermedia (Olivier and Odlaug, 1938) Odlaug, 1940; A. dubia Chandler and Rausch, 1946; A. minuta Chandler and Rausch, 1946, A. taxidea Swanson and Erickson, 1946; A. canadensis Webster and Wolfgang, 1956. Hosts: Procyon /o/or (Linn.). Location: Small intestine. Localities: Bayou Duplantier (East Ba- ton Rouge Par.); 2 mi. N. of Hwy. 190 on Atchafalaya River (Pointe Coupee Par.). Univ. Nebraska State Mus., Man- terLab. Coll. No. 21194 Diagnosis (based on ten mature speci- mens): Body indistinctly bisegmented, 1250 (1160-1420) long; forebody scoop- shaped, 680 (650-770) long by 565 (500- 580) wide, with pseudosuckers present on both sides of oral sucker; hindbody conical, 540 (460-650) long by 470 (450- 530) wide, containing reproductive or- gans. Forebody cuticle completely cov- ered with small spines; hindbody smooth. Oral sucker terminal, 86 (80-95) long by 81 (70-90) wide. Prepharynx pre- sent, short. Pharynx 66 (60-70) long by 48 (45-50) wide. Esophagus short, often not apparent. Ceca terminate near post- erior end of body. Acetabulum wider than long, 73 (70-80) long by 90 (85-93) wide. Holdfast broadly oval, variable in size, anterior extension partially or com- pletely covers the acetabulum. Testes two, not equal; anterior testis irregularly oval, laterally displaced on either side of midline, 188 (155-230) long by 206 (190- 230) wide; posterior testis dumbbell- shaped, much wider than anterior testis, 183 (150-220) long by 335 (355-400 wide, posterior testis with ventro-medial groove to allow passage of ceca and uterus. Seminal vesicle expanded into a muscular ejaculatory duct. Ovary reniform, located at forebody-hindbody juncture, 94 (90-100) long by 147 (135- 155) wide. Mehlis gland opposite anterior testis. Uterus makes an initial course into holdfast region and then pas- ses posteriorly emptying into the genital atrium. Genital atrium posterior, subterminal on dorsal surface. Vitellaria occupies much of the forebody from the acetabulum to anterior testis, never ob- served posterior to anterior testis. Vitel- line ducts unite to form a vitelline re- servoir at the level of anterior testis. Eggs large, few, 106 (102-1 10) long by 63 (60-65) wide. Excretory pore subtermi- nal on ventral side. Discussion: Babero (1960) tentatively listed Alaria taxidea as an intestinal in- habitant of the skunk. Mephitis mephitis, in Louisiana. Johnson (1979) relegated No. 2 Trematodes of Mammals 111 1 Figures 1-7. 1. Alaria mustelae, ventral view, from Procyon lotor.2. Pharyngostomoides procyonis , ventral view, from Procyon lotor. 3. Maritreminoides nettae, ventral view, from Procyon lotor. 4. Baschkirovitrema incrassatum, ventral view, from Lutra canadensis. 5. Gyrosoma singulare, ventral view from Procyon lotor. 6. Apophallus venustus, ventral view, from Procyon lotor. 7. Parametorchis complexus, ventral view, from Procyon lotor. Scale in micrometers. 112 Tulane Studies in Zoology and Botany Vol. 22 A. taxidea to a junior synonym of A. mustelae based on life history and host specificity for muteHds. He demon- strated that specimens of /4. mustelae dif- fered in size and morphology when re- covered from different mustelid hosts. He found no natural A. mustelae infection in the raccoons he examined, but 1000 mesocercariae fed to a single raccoon yielded 14 adults. Tliey were similar morphologically to those from mustelid infections, but were decidedly smaller. Bosma (1934) stated that she recovered only A. mustelae metacercariae from raccoons. This is the first report of natural infec- tions of adult Alaria mustelae in raccoons. We found two of thirty rac- coons infected. One infection consisted of fifteen worms whereas the second in- fection consisted of several hundred ma- ture adults in the small intestine. Our specimens are smaller (1. 160-1.420 mm) compared to the upper range (2.4 mm) of the species reported by Johnson (loc. cit.), but are larger than his specimens from the experimental infections in the raccoon (0.88-1.00 mm). Unfortunately Babero (loc. cit.) did not list measure- ments for his specimens from skunks and we were unsuccessful in gaining access to them. However, we concur that A. tax- idea is a synonym of y4. mustelae and that the low number of naturally infected rac- coons, small size of worms, and the low number of adults recovered in experi- mental infections of raccoons indicates a physiological specificity for mustelids. Of interest from an epidemiological standpoint, Beaver et al. (1977) reported a mesocercarial infection in a human in Louisiana. They believed the meso- cercariae members of the genus Alaria and traced the infection to the consump- tion of an undercooked raccoon. John- son (loc. cit.) has proven experimen- tally, and we have found in nature, that raccoons may serve as definitive hosts for A. mustelae. However, given the low specificity of A. mustelae for raccoons and the complex migration of all species of Alaria in the definitive host (gut to lungs and back to gut) some mesocercar- iae may lose their way and lodge in other tissues. We have evidence to this effect in other species of Alaria (to be pre- sented elsewhere). This indicates that the raccoon is not only a definitive but a paratenic host as well. Pharvngostomoides procyonis Hark- ema, 1942 (Figure 2) Synonyms: P. ovalis Chandler and Rausch, 1946. Hosts: Procyon lotor (Linn.) Location: Small intestine. Localities: St. Landry, Pointe Coupee, Iberville, St. Martin, East Baton Rouge, Ascension, and Livingston parishes. Univ. Nebraska State Mus., Manter Lab. Coll. No. 21195. Diagnosis (based on ten mature speci- mens): Body indistinctly bisegmented, 1 160 (1080-1250) long; forebody scoop- shaped, 580 (520-620) long by 570 (520- 630) wide, with pseudosuckers present on both sides of the oral sucker; hind- body conical, 580 (540-620) long by 506 (450-550) wide. Forebody cuticle comp- letely covered with small spines; hind- body smooth. Oral sucker terminal, 74 (70-75) long by 90 (80-95) wide. Prepharynx 10 (5-15) long. Pharynx muscular, 57 (50-60) long by 51 (45-55) wide. Esophagus 22 (20-25) long. Ceca extend just posterior to testes, never reaching posterior end of body. Ace- tabulum spherical, posterior to intestinal bifurcation, 77 (75-80) long by 78 (75-80) wide, artially or completely covered by holdfast organ. Holdfast well developed, elongate, broadly oval, may reach as far as pharynx. Testes equal, spherical or oval, opposite but often diagonal, 237 (2(X)-250) long by 200 (180-230) wide, located in hindbody dorsal to ceca. Semi- nal vesicle coiled, posterior to testes, ex- panded into a muscular ejaculatory pouch, that terminates in a genital cone. Genital atrium posterior, subterminal on dorsal side. Ovary reniform or trans- No. 2 Trematodes of Mammals 113 versely oval, anterior to testes, 90 (85- 100) long by 137 (125-160) wide. Uterus makes a short ascending loop into hold- fast region, turns posteriorly and unites with ejaculatory pouch. Vitellaria mainly in forebody, extending from acetabulum to the anterior margin of testes. Vitelline ducts unite just posterior to ovary to form the vitelline reservoir. Eggs few, large, 92 (90-95 ) long by 60 (58-62) wide. Excretory pore subterminal on ventral side. Discussion: Pharyngostomoides is a stenoxenic genus reported only from rac- coons. Harkema (1942) described P. procyonis from raccoons in North Carolina and Texas. P. ovalis was subse- quently described by Chandler and Rausch (1946) in Michigan and was later synonymized with P. procyonis by Dubois (1963). Harkema and Miller (1964) surveyed raccoons from North Carolina, South Carolina, Georgia, Florida, and Virginia and found P. pro- cyonis present in each state. Further, they noted that there appeared to be two forms present in the raccoon. Beckerdite et al. (1971) raised both of these forms to full species. They described P. adeno- cephala and separated it from P. procyonis on the basis of: 1) larger size; 2} shape (spathulate rather than scoop- shaped); 3) presence of pseudosucker glands; 4) absence of an ejaculatory pouch; and 5) morphology of larval stages. Based on our dignosis we conclude that our specimens belong to P. procyonis. We are not able to compare the morphology of the larval stages since the life history in Louisiana is unknown. We have found P. procyonis to be the most frequent parasite encountered in raccoons; 29 of 30 examined were in- fected with this worm. The only excep- tion was a raccoon collected near a resi- dential area of Baton Rouge. Infections ranged up to an estimated 5000 worms in one raccoon. Family MICROPHALLIDAE Travassos, 1920 Maritreminoides nettae (Gower, 1938) Rankin, 1939 (Figure 3) Synonyms: Maritrema nettae Gower, 1938; Pseudospelotrema nettae (Gower, 1938) Hunter and Vernberg, 1953. Hosts: Procyon lotor (Linn.). Location: Small intestine. Localities: Atchafalaya R.at Krotz Spr- ings (St. Landry Par.); 5 mi. N. of Butte La Rose on Atchafalaya R. (St. Martin Par.); 10 mi. S. of Ramah on East Atch- afalaya Protection Levee (Iberville Par.). Univ. Nebraska State Mus., Manter Lab. Coll. No. 21196 Diagnosis (based on ten mature speci- mens): Body small, pyriform, 407 (390- 420) long by 231 (220-270) wide. Cuticle completely covered with small spines. Oral sucker subterminal, 45 (37-50) long by 57 (55-61) wide. Prepharynx 25 (15- 33) long. Ceca short, thickwalled, bi- furcating anterior to cirrus sac and ex- tending toward edges of body; never ex- tending past anterior level of aceta- bulum, 130 (120-143) long. Acetabulum pre-equatorial, larger than oral sucker, 63 (57-66) long by 63 (61-68) wide. Testes spherical, opposite, equal in size, 71 (60-83) long by 70 (60-77) wide. Cir- rus sac crescent-shaped, transverse, pre- acetabular; walls of cirrus sac thick with well developed seminal vesicle, prostate, and ejaculatory duct; ejaculatory duct ex- tends into a slender, protrusible cirrus. Genital atrium opens sinistral to aceta- bulum; atrium armed with small, stout spines. Ovary irregularly oval, usually overlapping acetabulum, 71 (64-81) long by 79 (75-80) wide. Uterus consists of loops between acetabulum and posterior stem of excretory vesicle. Vitellaria form a right angle to testes with the sides being anterior and lateral to each testis, a few follicles extend posterior to testes but never reach end of body. Eggs small. 114 Tulane Studies in Zoology and Botany Vol. 22 numerous, 20 (20-22) long by 11 (10-11) wide. Excretory pore terminal; excre- tory vesicle V-shaped. Discussion: Rankin (1939) erected the genus Maritreminoides for those microphallids with a long, protrusible cirrus and antero-lateral uterine coils. He placed in his genus M. nettae and two metacercarial forms from fishes; M. ob- stipum (Van Cleave and Mueller, 1932) and M. medium (Van Cleave and Muel- ler, 1932). Etges (1953) regarded the genus Maritreminoides valid, but trans- ferred M. obstipum and M. medium into the genus Maritrema Nicoll, 1907 because they did not possess a protrusi- ble cirrus. Sogandares-Bernal (1965) surveyed crayfishes in Louisiana for parasites and reported Maritrema obstipum from Cambarellus shufeldti (Fa- xon, 1881) and Procambarus clarkii (Girard, 1852). The metacercaria figured by Sogandares-Bernal is morphologically identical with the adult worms we have found in raccoons except for lacking a protrusible cirrus and being ovigerous. Presumably, Sogandares- Bernal assigned the metacercariae to the genus Maritrema based on the absence of a protrusible cirrus. We believe the metacercariae found in crayfishes by Sogandares-Bernal (loc. cit. ) to be the larval stage of the adults we have collected from raccoons. The fact that a cirrus was not figured for the metacercaria could well have been be- cause the worms were immature. Func- tionally, a protruded cirrus in a metacer- carial stage, unless progenetic, would be of dubious value. The location of the infected crayfishes, as reported by So- gandares- Bernal, is in the same basin as are the infected racoons. We have observed crayfish remains in many rac- coons. Though Maritreminoides nettae was de- scribed from ducks in Michigan (Gower, 1938), Harkema and Miller (1964) re- ported it from raccoons in North Carolina, South Carolina, and Georgia. We reported the finding of M. nettae in the small intestine of seven of thirty rac- coons in Louisiana. We also report what we believe to be the second intermediate host, i.e. the crayfishes of Sogandares- Bernal (loc. cit.). If life histories of closely related forms are examined, the first intermediate host of M. nettae is most probably an amnicolid snail. Family ECHINOSTOMATIDAE (Looss, 1902) Poche, 1926. Baschkirovitrema incrassatum (Diesing, 1850) Skrjabin, 1944 (Figure 4) Synonyms: Dipostomum incrassatum (Diesing, 1850; Echinostomum incras- satum (Diesing, 1850) Stossich, 1891. Hosts: Lutra canadensis Schreber. Location: Small intestine. Locality: Lake Verret (Assumption Par.) Univ. Nebraska State Mus., Manter Lab.,Coll. No. 21197. Diagnosis (based on 14 mature speci- mens): Body elongate, slender, 27.5. (23-31) mm long. Maximum width at level of acetabulum, 1170 (1100-1300). Head color reniform, 546 (500-580) wide, bearing 27 spines; a double row of 4 corner spines on each side, 148 (140- 160) long by 42 (35-50) wide; six lateral spines on each side are found in a single row and are increasingly larger from ven- tral to dorsal side, the largest lateral spine, 121 (100-145) long by 26 (25-30) wide; 7 dorsal, uninterrupted spines, 132 (120-150) long by 25 (24-26) wide. Oral sucker 262 (220-280) long by 287 (250- 300) wide. Prepharynx 142 (130-150) long. Pharynx 240 (220-250) long by 170 (140-200) wide. Esophagus 690 (600- 800) long. Ceca two, terminating near posterior end of body. Acetabulum very prominent, muscular, located in anterior fifth of body, 1190 (1100-1400) long by 990 (9(X)-1050) wide. Testes tandem, elongate, oval, in anterior third of body; anterior testis 1190 (970-1450) long by 265 (200-320) wide; posterior testis 1090 (870-1400) long by 246 (200-310) wide. No. 2 Trematodes of Mammals 115 Vas efferens originate on antero-lateral side of testes and unite at posterior margin of the acetabulum ; cirrus sac dor- sal, not extending posterior to the acetabulum, 1150 (1100-1400) long by 500 (400-550) wide; cirrus long, slender, and unarmed. Ovary round, dextral to midline, 300 (240-350) long by 260 (250- 280) wide. Mehlis gland diffuse, imme- diately posterior to ovary. Uterus with short descending coil; ascending uterus forms transverse intercaecal loops be- tween Mehlis gland and acetabulum; dis- tal vessel of uterus forms a metraterm that opens at the common genital pore just anterior to acetabulum. Vitelline follicles extend from middle of anterior testis to posterior end of body, filling most of lateral space behind testes. Vit- elline ducts originate at middle of an- terior testis and unite to form a vitelline reservoir at level of Mehlis gland. Ex- cretory pore terminal. Eggs abundant, 109 (105-115) long by 64 (60-66) wide. Discussion: The geographical localities from which Baschkirovitrema incras- satum have been reported are of consid- erable interest. Braun (1901) described B. incrassatum from the otter, Pteronura brasiliensis Zimmerman, in South America; Beverly-Burton (1960) and Myers et al. (I960) report B. incrassatum from L. maculicollis Lichtenstein and Aonyx capensis (Schinz) from Africa; and Sawyer (1961), Harkema and Miller (1968), and Fleming et al. (1977) have identified B. incrassatum from L. canadensis in North America. Harkema and Miller (loc. cit.) re- ported B. incrassatum from North Carolina and stated that they compared their specimens with those of Sawyer (loc. cit. ) from Georgia and to specimens deposited in the National Museum by W. J. Hamilton from otters in New York. They reported all specimens from North America were similar and that they "compared favorably" with the meas- urements given by Beverly-Burton (loc. cit.) from Africa. The present report is the first to present measurements from North American material. We have found that our specimens from Louisi- ana are much larger than those reported from South America (Braun, 1901) and from Africa (Beverly-Burton, 1960) and with no apparent overlap (Table 1). In addition, the oral sucker and acetabulum of our specimens are proportionately larger than previously described. As indicated by the measurements, our specimens are more closely aligned with the material from South America than those from Africa. Since life histories of any of these forms are enig- matic, it is difficult to ascribe differences at this time to intra- or interspecific variation. Family PSILOSTOMIDAE Looss, 1900 Gyrosoma singulare Byrd, Bogitsh, and Maples, 1961 (Figure 5) Synonyms: Originally described as Grysoma singularis Byrd et al., but Yamaguti (1971) emended the genus to Gyrosoma. Hosts: Procyon lotor (Linn.). Location: Small intestine. Localities: Bayou Duplantier and Spanish Lake (East Baton Rouge Par.); Atchafalaya R. between Keith Lake and Krotz Springs (St. Landry Par.); and Head of Island (Livingston Par.). Univ. Nebraska State Mus., Manter Lab. Coll. No. 21198. Diagnosis (based on ten mature speci- mens): Body lanceolate, relatively thick, 632 (580-680) long; widest at level of acetabulum, 222 (200-265) wide. Cuticle thick, beset with short, stout spines; spines set in dense rows anteriorly and become less numerous behind the post- erior testis. Oral sucker subterminal, 55 (52-59) long by 56 (52-59) wide. Pre- pharynx 15 (13-17) long. Pharynx barrel- shaped, 35 (30-41) long by 36 (33-39) wide. Esophagus distinct, 14 (22-41) long. Ceca bifurcating anterior to com- mon genital pore, diverge laterally to acetabulum and terminate midway bet- ween posterior testis and caudal end of body. Acetabulum prominent, pre- 116 Tulane Studies in Zoology and Botany Vol. 22 Table 1. Comparative measurements of Baschkirovitrema incrassatum from Africa, South America, and North America. Africa South America North America Beverley-Burton (1960) Braun(1901) 7-19 mm (present study) Body length 12.50-16.55 mm 27.5 (23-31) mm No. of head spines 27 27 27 Comer spines 108-154X28-37 104X31 148 (140- 160) X 42 (35-50) Lateral spines 105-130X28-32 - 121 100-145) X 26 (25-30) Dorsal spines 109-144X30-34 83-93X21 132 (120-150) X 25 (24-26) Oral sucker: length 190-220 166-250 262 (220-280) Oral sucker: diameter 220-260 187-208 287 (250-300) Acetabulum: length 870-1170 - 1190(1100-1400) Acetabulum: diameter 810-950 1000 990(900-1050) Pharynx: diameter 120-210 73-83 170(140-200) Testes: length 1120-1500 1000 1140(870-1450) Testes: diameter 320-480 400 256 (200-320) Ovary: length 320-390 330 300 (240-350) Ovary: diameter 320-370 266 260 (250-280) Eggs 108-123X54-62 104X73 109 (10^-115) X 64 (60-66) equatorial, 89 (81-92) long by 82 (70-92) wide. Testes oval, diagonal (rarely tan- dem), contiguous and often overlapping; anterior testis 76 (66-88) long by 85 (81- %) wide; posterior testis 86 (72-%) long by 83 (77-94) wide. Cirrus sac promi- nent, dorsal to acetabulum, containing seminal vesicle, well developed prostate, and a short, muscular cirrus. Genital pore midventral between cecal bifurca- tion and acetabvulum; a well developed sphincter muscle is observed when the genital pore is constricted. Ovary ovoid, lying immediately postero-dorsal to acetabulum, 45 (42-48) long by 42 (40- 44) wide. Uterus short, ascends directly to genital pore; usually only one egg ob- served in the uterus at one time, but occasionally two. Egg thick-shelled, large in comparison with body size, 105 (100-110) long by 64 (60-75) wide. Vitel- laria extend laterally from the genital pore to caudal end of body; each follicle composed of large, clearly defined vitel- line cells. Vitelline ducts extend medially between testes and acetabulum to form a large vitelline reservoir. Excretory sys- tem composed of a terminal excretory pore; a Y-shaped excretory vesicle that bifurcates at caudal end of ceca; and arms that extend anteriorly to level of the pharynx. Discussion: The genus Gyrosoma was created by Byrd et al. (1961) to accom- modate specimens collected from a single raccoon in Georgia. They con- sidered these worms to be distinct from other psilostomids based on: 1) the diagonal arrangement of the testes; 2) triangular shape of the testes; 3) a short uterus; and 4) a single, large egg in the uterus. To our knowledge, the only other mention in the literature of G. singulare is that of Harkema and Miller (1964) who state that they have also found it in Georgia and South Carolina. We have collected G. singulare from the small intestine of 24 raccoons in Lx)uisiana. Infection ranged from a few to several hundred worms. This appar- No. 2 Trematodes of Mammals 117 ently stenoxenic species is most easily distinguished by the presence of a large egg which may approach one fourth the size of the worm. No morphological data have been reported for this species other than that contained in the original de- scription. Our measurements signi- ficantly extend the size range of this species (580-680 compared to 290-480) and we find that our specimens have more spherical shaped testes than the triangular or half-mooned shape re- jKDrted by Byrd et al. (loc. cit.). Family HETEROPHYIDAE (Leiper, 1909) Odhner, 1914 Apophallus venustus (Ransom, 1920) Cameron, 1936 (Figure 6) Synonyms: Cotylophallus venustus Ransom, 1920; C. similis Ransom, 1920; Rossicotrema venustum Ciurea, 1924. Hosts: Procyon lotor (Linn.). Location: Small intestine. Locality: 10 mi. S of Ramah on the East Atchafalaya Protection Levee (Iberville Par.). Univ. Nebraska State Mus., Manter L2b. Coll. No. 21199 Diagnosis (based on ten mature speci- mens): Body elongate, oval with a slight constriction between acetabulum and ovary, 1018 (820-1150) long by 563 (450- 650) wide. Cuticle with small spines over anterior two thirds; posterior devoid of spination. Oral sucker unomamented, subterminal, 56 (46-66)long by 70 (55- 79) wide. Prepharynx distinct, 13 (11-24) long. Pharynx 54 (50-57) long by 52 (48- 55) wide. Esophagus long and slender, 123 (99-132) long. Ceca extend to caudal end of body. Acetabulum small, weakly defined, located in genital sinus posterior to genital openings, 41 (31-44) long by 41 (39-44) wide. Testes oblique, globular or ovoid; anterior testis 129 (99-154) long by 163 (99-187) wide; posterior testis 133 (110-165) long by 180 (150-224) wide. Vas efferens unite at level of ovary to form a conspicuous sigmoid seminal ves- icle. Cirrus absent. Ovary ovoid, dextral to midline, 104 (99-115) long by 110 (99- 125) wide. Seminal receptacle large, transversely oval, and lying directly post- erior to ovary. Uterus makes 3-4 trans- verse, intercaecal loops before entering genital sinus. Vitellaria unite across body between the intestinal bifurcation and acetabulum, then extend laterally to posterior end of body. Vitelline ducts unite to form a vitelline reservoir bet- ween seminal vesicle and testes. Eggs 35 (33-37) long by 20 (17-22) wide. Excret- ory pore terminal; posterior stem of ex- cretory vesicle sigmoid as it courses bet- ween the testes. Discussion: Apophallus venustus has, heretofore, been reported from eastern coastal states and provinces of North America (Cameron, 1936; Babero and Shepperson, 1958; and Harkema and Miller, 1964). Yamaguti (1971) consid- ers ^4. venustus conspecific with its Euro- pean relative A. donicus (Skrjabin and Lindrop, 1919) Price, 1931, but we pre- fer to follow Cameron (loc. cit.) and rec- ognize A. venustus as a distinct species based on its greater anterior extent of vitellaria and its larger size. The life history of A. venustus was studied by Cameron (1937, a) and con- sists of an op>erculate snail, Goniobasis livescens (Menke), as first intermediate host, a wide variety of fishes as second intermediate host, and any fish-eating mammal as the definitive host. Cameron (1937,b) reported a human infection in Canada with A. venustus eggs being found in the feces. The epidemiology was traced to the presumed ingestion of fish. The presence of A. venustus in Louisiana is of epidemiological interest since many of its people subsist, to a large degree, off the land. Small infec- tions may p>erhaps go unnoticed because adequate diets prevent loss of vigor. However, Africa et al. (1935), working in the Philippines, found a high correla- tion between cardiac alteration and heterophyid infection. Kean and Breslau (1964) state that a large number of cardiac fatalities (14.6%) in the Philip- 118 Tulane Studies in Zoology and Botany Vol. 22 pines are attributed to heterophyid eggs which find their way into the circulation. These reports, coupled with the already mentioned human infection with A. ven- ustus should alert epidemiologists and pathologists to the presence of this speies and its possible involvement inhuman in- fection in Louisiana. We found four of thirty raccoons in- fected with A. venustus. The infections were located throughout the small in- testine and not localized only in the je- junum and ileum as reported by Came- ron ri937,a). Family OPISTHORCHIIDAE Braun, 1901 Parametorchis complexus (Stiles and Hassal, 1894) Skrjabin, 1913 (Figure 7) Synonyms: Ditoma complexum Stiles and Hassal, 1894. Hosts: Procyon lotor (Linn.). Location: Biliary system. Localities: Sherburne (Pointe Coupee Par.); 10 mi. S. of Ramah (Iberville Par.). Univ. Nebraska State Mus., Manter Lab. Coll. No. Diagnosis (based on ten mature speci- mens): Body flat, elongate, anterior end attenuate and posterior rounded, 7.8 (6.0-10.5) mm long by 2.2 (1.6-2.9) mm wide. Entire cuticle spinous; spines less dense posterior to testes. Oral sucker terminal, 272 (230-320) long by 326 (330- 400) wide. No prepharynx. Pharynx muscular, 253 (210-290) long by 233 (220-270) wide. Esophagus short, 113 (70-150) long. Ceca large, coursing sinu- ously around organs until reaching post- erior end of body. Acetabulum weak, inconspicuous, located in anterior one fourth of body, 286 (260-330) long by 305 (280-330) wide. Testes tandem, lobate, located in posterior half of body; an- terior testis 913 (670-1250) long by 956 (800-1150) wide; posterior testis usually larger and possessing more lobes, 994 (800-1400) long by 955 (730-1200) wide. Vas deferens enlarged to form seminal vesicle that courses anteriorly, dorsal to uterus, and empties into genital pore. Genital pore located ventrally, midway between acetabulum and medially positioned vitellaria. Ovary trilobed, pretesticular, 405 (300-500) long by 494 (390-610) wide. Seminal receptacle vari- able in shape, usually larger than ovary, lying immediately posterior to, and often overlaping, the ovary. Uterus volumin- ous, with tightly packed loops loosely bordered by the ceca, occupying most of the space between ovary and aceta- bulum. Vitellaria laterally displaced in anterior half of body; unite medially be- twen intestinal bifurcation and genital pore. Vitelline ducts unite ventral to ov- ary to form a vitelline reservoir. Eggs oval, some constricted anteriorly, 28 (26- 30) long by 16 (15-17) wide. Excretory pore terminal; posterior stem of excret- ory vesicle sigmoid as it passes between testes, eventually giving rise to a Y-shaped excretory vesicle. Discussion: Stiles and Hassal (1894) originally described P. complexus from specimens found inthe biliary system of cats in Maryland and New York. Since that time, additional species have been described from the biliary system of carnivores: P. noveboracensis Hung, 1926 from the cat; P. intermedius Price, 1929 from the fox, Vulpes fulva (Desmarest); P. canadensis Price, 1929 from Mustela vison ; and P. manitobensis Allen and Wardle, 1934 from the dog. Cameron (1944) examined the morphol- ogy of these worms and concluded that the latter four species were synonyms of Metorchis conjunctus (Cobbold, 1860) Looss, 1899. He based his separation on the anterior disposition of the vitellaria. In M. conjunctus the vitellaria are later- ally displaced while the genus Para- metorchis is characterized by the anterior union of the vitellaria. Came- ron's proposal left Parametorchis a monotypic genus. Yamaguti (1971) incorrectly stated that Cameron synon- ymized P. intermedius, P. canadensis, P. noveboracensis, and P. manitobensis with P. complexus. No. 2 Trematodes of Mammals 119 Harkema and Miller (1964) reported P. complexus from one of 320 raccoons they examined in the southeastern Un- ited States (the infected raccoon was found in North Carolina). We have found P. complexus in five raccoons in Louisiana. The worms inhabit all levels of the biliary system One raccoon har- bored 91 mature P. complexus and the bile duct exhibited marked hypertrophy. SUMMARY TTie following trematodes were col- lected from small mammals in southern Louisiana: Alaria mustelae Bosma, 1931 ; Fibricola cratera (Barker and Noll, 1915) Dubois, 1932; F. lucida (La Rue and Bosma, 1927) Dubois and Rausch, 1950; Pharyngostomoides procyonis Har- kema, 1942; Heterobilharzia americana Price, 1929; Apophallus venustus (Ransom, 1920) Cameron, 1936; Para- metorchis complexus (Stiles and Hassal, 1894) Skrjabin, 1913; Baschkirovitrema incrassatum (Diesing, 1850) Skrjabin, 1944; Brachylaima virginianum Dicker- son, 1930; Gyrosoma singulare Byrd, Bogitsh, and Maples, 1%1; Hasstilesia texensis Chandler, 1929; Maritremi- noides nettae (Gower, 1938) Rankin, 1939; and Rhopalias macracanthus Chandler, 1932. Trematodes reported from. Louisiana for the first time are: Alaria mustelae, Pharyngostomoides procyonis, Mari- treminoides nettae, Baschkirovitrema in- crassatum, Gyrosoma singulare, Apo- phallus venustus, and Parametorchis complexus. The raccoon is a new definitive host for A. mustelae. A. mustelae shows a dis- tinct specificity for mustelids rather than procyonids based on larger size of worms, low number of naturally infected raccoons, and the low number of adults recovered from experimentally infected raccoons. During migration of larval A. mustelae in the raccoon, mesocerariae may lose their way and lodge in other tissues. The raccoon is, then, both a de- finitive and paratenic host. This finding becomes important in light of the recent report of Alaria mesocerariae in man in Louisiana (Beaver et al., 1977). Pharyngostomoides procyonis is the most frequently encountered trematode in raccoons in Louisiana. Though it has been reported from the eastern coastal states and Texas, it likely is found throughout the other Gulf Coast states as well. Metacercariae reported in crayfishes as Maritrema obstipum by Sogandares- Bemal (1965) are identical to adult Maritreminoides nettae recovered from raccoons, except for a protrusible cirrus and being ovigerous. The infected crayfishes are found in the same river basin as are the infected raccoons. We propose that they are the same species based on morphological, geographical and ecological criteria. Specimens identified as Baschkirovi- trema incrassatum have been reported in otters from North America, South America, and Africa. Since speciation in otters has ocurred, these worms possibly may be in the process of pursuing distinct evolutionary pathways. Our specimens from North America are much larger than previously ref)orted and with no ap- parent overlap. Based on reported mea- surement, our specimens notably are more closely aligned with the material from South America than from Africa. The presence of A. venustus in Louisiana is of epidemiological signifi- cance since a human infection involving this species has been previously re- ported. The metacercariae of A. venus- tus is foundin fish and all fish-eating mammals are susceptible, including hu- mans. Since other heterophyids have been found in be the cause of cardiac fatalities, both epidemiologists and pathologists should be aware of its pre- sence and the possible involvement in human infection in Louisiana. 120 Tulane Studies in Zoology and Botany Vol. 22 LITERATURE CITED AFRICA, C. M., GARCIA, E. Y., and DE LEON, W. 1935. Intestinal Heterophyidae with cardiac involvement. A contribution to the etiology of heart failure. Philippine J. Public Health 2: 1-22. BABERO, B. B. 1960. A survey of parasitism in skunks (Mephitis mephitis) in Louisiana, with observations on pathological damages due to helminthiasis. J. Parasitol. 46: 26-27. , and SHEPPERSON, J. R. 1958. Some helminths of raccoons in Georgia. J. 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Morphology and life history of Alaria mustelae Bosma, 1931 (Tre- matoda: Diplostomatidae) from Minnesota mustelids. J. Parasitol. 65: 154-160. KAPLAN, E. H. 1964. Heterobilharzia americana Price, 1929, in the opossum from Louisiana. J. Parasitol. 50: 797. KEAN, B. H., and BRESLAU, R. C. 1964. Para- sites of the Human Heart, Chapter 10. Cardiac Heterophyidiasis. Grune & Stratton, New York. pp. 95-103. LUMSDEN, R. D., and ZISCHKE, J. A. 1961. Seven trematodes from small mammals in Louisiana. Tulane Stud. Zool. and Bot. 9: 87- 98. MALEK, E. A., ASH, L. R., LEE, H. F., and LITTLE, M. D. 1961. Heterobilharzia infec- tion in the dog and other mammals in Louisiana. Parasitol. 47: 619-623. MYERS, B.J., WOLFGANG, W., KUNTZ, R. E. 1960. Helminth parasites from vertebrates taken in the Sudan (East Africa). Can. J. Zool. 38: 833-836. RANKIN, J. S. 1939. Studies on the trematode family Microphallidae, Travassos, 1921. III. The genus Maritrema Nicoll, 1907, with de- scription of a new species and a new genus, Maritreminoides. Am. Midi. Nat 22: 438-451. SAWYER, T.K. 1%1. The American Otter, Lutra canadensis vaga, as a host for two species of trematodes previously unreported from North America. Proc. Helminthol. Soc. Wash. 28: 175-176. No. 2 Trematodes of Mammals 121 SOGANDARES-BERNAL. F. 1965. Parasites from Louisiana crayfishes. Tuiane Stud. Zool. and Bot. 12: 79-85. STILES, C. W., and HASSAL, A. 1894. A new species of fluke (Distoma (Dicrocoelium) com- plexum) found in cats in the United States, with bibliographies and diagnosis of allied forms. Vet. Mag. 1:413-432. YAMAGUTI, S. 1971. Synopsis of Digenetic Tre- matodes of Vertebrates. Vo. I. Keigaku Publ. Co., Ltd., Tokyo. 1,100 p. May 3, 1981 TULANE STUDIES IN ZOOLOGY AND BOTANY VOLUME 22 INDEX TO SCIENTIFIC NAMES (New taxonomic entities in lioldface) Acacia, 1, 4, 8 Acer negundo, 19 saccarinum, 19 Agave, 1,2,8, 12 Agelaius phoeniceus, 43 Alaria canadensis, 110 dubia, 110 freundi, 110 intermedia, 110 marcianae, 109 minuta, 110 mustelae, 109-112 /ox/rf^fl, 110-111 Alburnops, 67 y4me/va chrysolaema, 2 Ammocrypta vivax, 70 Amorpha fruticosa, 19 Aonyx capensis, 115 Apophallus donicus, 117 venustus, 111,117-118 Ardea herodias, 44 Asclepias syrica, 19 Barbarea vulgaris, 19 Baschkirovtrema incrassatum. 111, 114-115 B^ru/a, 29 «/gra, 19 Bothriechis lansbergii, 87 Bothriopsis proboscideus, 86-87 Bothrops, 86 brachystoma, 86-87 godmani, 102 lansbergi, 85-87, 102 rto^M/a, 85-107 nummifera, 102 yucatanica, 88 Brachylaima virginianum, 109-111 Bromus tectorum, 19 Cambarellus shufeldti, 1 14 Carex sp., 19 Ceratophyllum sp. , 20 Chelydra serpentina, 31, 41 Chrysemys, 5 1 , 54 p/cw. 27-28, 30,34,41,52 p. fee//., 43 Citrobacter freundi, 31 Corkum, Kenneth C. article, 109-121 Comus obliqua, 19 Corvu5 brachyrhynchos, 43 Cotylophallus similis, 117 venustus, 117 Cycloloma atriplicifolium, 19 Didelphis virginiana, 109 Dipostomum incrassatum, 114 Ditoma complexum, 118 Echinostomum incrassatum, 1 14 Elodea sp., 19 Emydoidea blandingi, 41 concinna, 61 floridana, 61 hieroglyphica, 61 mobilensis, 61 orbicularis, 61 Epicrates f. fordi, 12 Equisetum arvense, 19 Erigeron annus, 19 Fahey, Kenneth Mark, article, 49-66 Fibricola cratera, 109-110 /Mc/rfa. 109-110 Fraxinus pennsylvanica, \a.x. subinterrima, 19 Galium obtusum, 19 Gonibasis livescens, 117 Gopherus agassizi, 34 Graptemys, 17-20, 24, 26-27, 30, 34, 37, 39, 42-43, 46 geographica, 22, 24, 27, 35, 39-42, 45 ouachitensis, 17-18, 20, 22, 4-27, 30-42, 44-45 o. ouachitensis, 17 o. sabinensis, 17-18 pseudogeographica, 17, 20, 22, 24-27, 30 42, 44-45 p. kohni, 17,31 p. pseudogeographica, 17-18 pM/c/jra, 23-25, 34, 40 Gyrosoma, 117 singulare, 111,115-117 Hasstilesia texensis, 109-110 Heins, David C, article, 67-84 Heterobilharzia americana, 1 10 He'eranthera dubia 19 La,.hesis brachystoma, 87 Laportea canadensis, 19 Larus delawarensis, 43 Lemna, 40 minor, 19 Lepidium densiflorum, 19 Lutra canadensis, 43, 109, 114-115 maculicollis, 115 Lynchnis alba, 19 Maritrema, 114 INDEX TO AUTHORS AND SCIENTIFIC NAMES — Continued Maritreminoides, 1 14 medium, 114 nettae, 111,113-114 obstipum, 114 McCranie, James L., article, 85-109 Mephitis mephitis, 109-110 Metorchis conjunctus, 118 Mollugo verticillata, 19 Mustela vison, 109-118 Myriophyllum sp., 20 Malaclemys terrapin, 31 Nelumbo lutea, 19 Notropis, 81 longirostris, 67, 81-82 sabinae, 67-84 venustus, 70 Nymphaea tuberosa, 19 Oenothera biennis, 19 Parametorchis canadensis, 118 complexus, 111,118-119 intermedins, 118 manitobensis, 118 novabracensis, 118 Parthenocissus inserta, 19 Phalaris arundinacea, 19 Pharynogostomoides ovalis. 112-113 procyonis, 111-113 Phragmites communis, 19 Phyllodactylus, 12 Placobdella parasitica, 17, 43, 46 Polygonum convolus, 19 Populas deltoides, 19 Porras, Louis, article, 85-107 Porthridium, 86-102 nasutum, 86-87 «. nasutum, 87 n. sutum, 87 nasutus, 87 Potamogeton, 40, 42 americanus, 19 crispus, 19 foliosus, 19 nodosus, 19 Procambarus clarki, 1 14 Procyon lotor, 43, 109-113, 115, 117-118 Pseudem v.v concinna, 49-64 hieroglyphica, 49-52 mobilensis, 49-52 suwanniensis, 50-54, 61-62 texana, 50-5 1 , 54 floridana. 49-64 /lov/, 49-52 peninsularis, 50-51, 53, 61-62 nelsoni. 54-55 rubriventris, 34, 55 5cnpra. 24, 27, 30, 34, 42, 52, 55 elegans, 34 Pseudospelotrema nettae, 113 Pteroura brasiliensis. 115 Quiscahis quiscala. 17, 43, 46 Rhopalias macracanthus. 109 /?/m5 radicans, 19 Rossicotrema venustum, 117 5fl//.r sp., 19 fluviatilis, 19 officinalis, 19 Sagittaria latifolia, 19 Schwartz, Albert, article, 1-15 Scirpus validus, 19 Shoop, Wesley L, article, 109-121 Solanum dulcemera, 19 Sparganium americanum. 19 Sphaerodactylus, 1,3-6, 9, 12, 14 altavelensis, 2, 113 asterulus, 1,5, 9-14 cinereus, 2 cryphius, 13 difficilis, 13-14 diffici I is complex, 12-13 leucaster, 1-9, 11-15 notatus group, 13 ocoae, 7, 13 rhabdotus, 1-6,9,12-15 5/jrevc/. 1-5,7-9, 11, 13-14 s/ir^ve/ group, 12, 14-15 Spirodella polyrhiza. 19 Sporobolus cryptandrus, 19 Svh'ilagus aquaticus, 109 Terrapene Carolina, 27 ornata, 27, 34 Testudo concinna, 50, 61 floridana, 50, 61 Thanatophis, 86 5M/as. 86-87 Thanatos, 86 5ufu5, 86-87 Tradescantia ohiensis, 19 Trimeresurus brachystoma, 87 nasutus, 87 yura/an/cu.s, 88 7"r/oAiyj: mulicus, 43 spiniferus, 28, 43 Ulmus americana, 19 ^Urocyon cinereoargenteus, 109 INDEX TO AUTHORS AND SCIENTIFIC NAMES — Continued Urtica dioica, 19 Vallisneria, 40, 42 Verbascum thopsus, 19 Vitis riparis, 19 Vogt, Richard C, article, 17^ Vulpesfulva, 43, 118 Wilson, Larry David, article 85-107 Wolfiella columbiana, 20 Zizania aquatica, 19 cAcme Bookbindins Co.. inc. 100 CambriilSi $). 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