North American Recent Soft-Shelled Turtles (family Trionychidae)

Volume 13, No. 10, Pp. 429-611, Pls. 31-54, 24 Figs.

Published February 16, 1962

UNIVERSITY OF KANSAS

Lawrence, Kansas

PRINTED IN

THE STATE PRINTING PLANT

TOPEKA, KANSAS

1962

28-7818

North American Recent Soft-shelled Turtles

(Family Trionychidae)

ROBERT G. WEBB

CONTENTS

PAGE

CONTENTS 431

INTRODUCTION 433 Collecting Methods 434 Materials and Procedure 437 Acknowledgments 439

TAXONOMY 439 Family Trionychidae Bell, 1828 439 Genus Trionyx Geoffroy, 1809 443 Variation 445 Secondary Sexual Variation 446 Ontogenetic Variation 449 Geographic Variation 453 Character Analysis 460 Composition of the Genus Trionyx in North America 476 Artificial Key to North American Species and Subspecies of the Genus Trionyx 476 Systematic Account of Species and Subspecies 479 Trionyx ferox 479 Trionyx spinifer 486 Trionyx spinifer spinifer 489 Trionyx spinifer hartwegi 497 Trionyx spinifer asper 502 Trionyx spinifer emoryi 510 Trionyx spinifer guadalupensis 517 Trionyx spinifer pallidus 522 Trionyx ater 528 Trionyx muticus 531 Trionyx muticus muticus 534 Trionyx muticus calvatus 539

NATURAL HISTORY 541 Habitat 541 Daily and Seasonal Activity 547 Diurnal Habits 547 Behavior Adaptations 549 Movement 552 Nocturnal Habits 553 Seasonal Occurrence 553 Food Habits 555 Reproduction 558 Size of Males at Sexual Maturity 558 Size of Females at Sexual Maturity 560 Sexual Activity 563 Deposition of Eggs 565 Reproductive Potential 568 Eggs 572 Incubation and Hatching 573 Age and Growth 574 Mortality 576 Parasites 576 Economic Importance 577

EVOLUTIONARY HISTORY 578 Distribution 578 Relationships 579 Fossils 582 Phylogeny 585 The Importance of the Study of Turtle Populations in Relation to the History of River Systems 588

SUMMARY 590

LITERATURE CITED 594

INTRODUCTION

Is it true that the greater the degree of resemblance between two populations the shorter the time the two have been spatially isolated? Are aquatic environments more stable than terrestrial environments? These questions occurred to me while I was collecting turtles from river systems of the Gulf Coast. As a general rule, each kind of turtle seemed to occur throughout one continuous river system or large tributary, and with no barriers to dispersal therein and with the lapse of enough time for a population to reach its limits of dispersal, the question arose, "Where do subspecies and zones of intergradation occur?" It seemed logical to think that each isolated and continuous aquatic environment would not contain more than one subspecies of the same species. In terrestrial environments subspecies and transitions between them were recognizable. Terrestrial habitats were continuous for longer distances than the isolated, aquatic habitats. But, different species of turtles prefer different kinds of aquatic habitats. Also, barriers occur in large drainage systems, such as the Mississippi, where, in general, the western tributaries are sluggish, turbid and shallow, and the eastern tributaries are fast-flowing, clear and deep. But in young, relatively small, river systems that do not traverse radically different physiographic regions, and that show no gross ecological differences, habitats or microhabitats that do exist probably are only partial barriers and seem not to prevent the dispersal of most kinds of aquatic turtles. Consequently, it seemed that study of the degree of difference between closely related populations of turtles that occurred in one drainage system, or in adjacent drainage systems would indicate the length of time, respectively, that the drainage system had been continuous or the length of time that two or more systems had been isolated from one another.

Rivers or series of river systems having endemic kinds of turtles or having the most kinds of turtles that are different from those in adjacent rivers may be the oldest geologically, or may have been isolated the longest. Knowledge of the kinds of turtles and their relationships and distribution could indicate chronological changes in aquatic habitats. Of course, modifying factors such as differences between populations of turtles in rates of evolutionary change, degrees of vagility, rates of dispersal, and overland migrations need to be taken into account.

My accumulation of data on soft-shelled turtles was begun in the early nineteen-fifties. Although American softshells have been discussed in a revisionary manner by Agassiz (1857), Siebenrock (1924), Stejneger (1944) and Neill (1951), the relationships of all the component populations have not hitherto been appreciated. The present account attempts to combine in one publication what is known concerning the taxonomy, geographic distribution, life history, and relationships of the Recent American species and subspecies of the genus Trionyx.

Collecting Methods

Nocturnal collecting, by hand, from a boat that was nosed among brush piles along the shore line of rivers (Chaney and Smith, 1950:323) in the early 1950's on rivers of the Gulf Coast drainage east of Texas yielded many turtles of the genus Graptemys but few softshells. Chaney and Smith (loc. cit.) reported only one softshell among 336 turtles taken in 21 collecting hours on July 5, 6 and 7 on the Sabine River; Cagle and Chaney (1950:385), however, recorded 11.6 per cent softshells of 208 turtles (collecting time not stated) taken on the Caddo Lake Spillway in Louisiana. Using hoop-nets is probably the most efficient method for collecting softshells considering the time and effort involved, and is the chief method I have used. Lagler (1943a:24) mentioned the use of watermelon rind as an effective bait. Kenneth Shain (field notes) trapped T. spinifer emoryi in hoop-nets baited with bread. I have used chopped fresh fish with most success; canned sardines have also been satisfactory. These baits seem to be more successful for trapping spinifer than they are for muticus. Hoop-nets were used to trap turtles in Lake Texoma, Oklahoma, from June 14 to July 2, 1954. The number of traps (usually four, rarely five) and trapping success varied with location. Of 156 turtles, 19 (12%) were T. spinifer and one was T. muticus.

Trotlines and set lines frequently catch softshells; sport fishermen often complain of catching these turtles on hook and line. Live worms, soft-bodied insects, small crawfish, minnows, small pieces of fish and other kinds of meat are adequate bait. Capture depends on the skill of attachment of the bait and the size of hook used. In my experience, softshells (mostly spinifer) were taken on trotlines that were set in lakes or the slower-moving parts of rivers a few inches below the surface. I have records of only two muticus taken on trotlines. Goin (1948:304) stated that commercial fishermen catch softshells on trotlines set for catfish on the bottom of river beds. Evermann and Clark (1920:595) found softshells to be caught more often than any other kind of turtle in traps, on set lines, and by anglers in Lake Maxinkuckee, Indiana. Some residents of the South tell of so placing baits that turtles are lured to tread water against an object set with recurved hooks upon which the webbing of the forelimbs are impaled.

Individuals of muticus and spinifer frequently bury themselves in sand in shallow water and can be collected by hand by noting swirls or disturbances on the bottom caused by a turtle withdrawing its head (Conant, 1951:156, 159). Professional turtle collectors take them by "noodeling" (Conant, op. cit.:160); Lagler (1943a:22) elaborated on the method of "noodling." P. W. Smith (1947:39) remarked that 20 or more softshells were taken "within a few hours by probing sand bars at the water edge" near Charleston, Illinois. From a distance I observed an individual of T. s. asper bury itself in shallow water on the Escambia River, Florida. Small individuals of muticus have been taken by hand along the shore of Lake Texoma. Along the Flint River near Bainbridge, Georgia, two hatchlings that were buried in sand in shallow water emerged at my approach and scurried a few inches, then buried themselves again. Larger turtles seem to be more wary. One that was disturbed, emerged from the sand and swam toward deep water.

In clear water, water-goggling may be effective in securing softshells. Marchand (in Carr, 1952:417-18) mentioned that softshells (ferox) can be found buried in deep water with only the heads visible; the turtles are not easily frightened under water and may be captured by grasping their necks. A similar technique described by Allen and Neill (1950:3) resulted in the capture of trionychid turtles. In clear water of the White River, Arkansas, I collected a few softshells by hand as they lay on the bottom.

In shallow-water areas of large rivers, lakes and tributaries, seining often procures softshells. Methods used in fisheries investigations such as the application of rotenone and electric shockers, and even dynamiting, sometimes yield soft-shelled turtles. Carr (1952:419) wrote that numbers of ferox were incapacitated by rotenone in Florida lakes, although no other species of turtle was affected. I captured a snapping turtle (Chelydra serpentina) that was immobilized by the current from an electric shocker in a small, alga-choked tributary of Cache Creek, Comanche County, Oklahoma; presumably turtles must come in close contact with the electrodes to be affected (see discussion by Gunning and Lewis, 1957:52).

The effectiveness of gill nets in trapping turtles is indicated by information kindly supplied by Mr. Alfred Houser on gill-net operations from July through December, 1952, under the direction of Mr. "Bud" Oldham, a commercial fisherman. The 4-inch mesh nets were in Lake Texoma at the mouth of Briar Creek, two miles south of Powell, Marshall County, Oklahoma, in 25 to 30 feet of water. Eighty to 90 per cent of the turtles secured were softshells; more were taken near shoreline than away from shore even though the depth was about the same. An average of only one turtle every four days was taken in July and August when the turtles presumably are most active (Table 1). One gill-net day is equivalent to one gill net, 200 yards long, operated for 24 hours.

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