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TABLE 1. Experimental parameters and associated burrow architectures. Terrarium sizes are in gallons. Sediment indicates the relative percentage of O = organic (coconut fiber), S = soil (clay loam), and Sa = fine- to medium-grained carbonate sand. Depth is the total depth of the sediment in centimeters. % Moisture is the mean moisture content of the sediment. Duration is split into two categories: expected time to completion of trial and the actual trial duration (in parentheses). Key to burrow architectures and modifications: V = vertical burrow, SV = subvertical burrow, H = helical burrow, J = J-shaped burrow, O = O-shaped burrow (T1=Type 1, T2=Type 2), c = chamber(s), b = branch(es), m = multiple entrances.

 

Experiment 1: Basic Morphology

Species

Specimens

Terrarium

Sediment

Depth

% Moisture

Duration

Burrow #s

Burrow Architectures

N. americanus

5

30

50/50 (O/S)

20

37%

7 (12)

NA26 A-B

SV, O (T1)

N. americanus

5

65

50/50 (O/S)

45

32%

7 (12)

NA23

SV

N. americanus

1

30

50/50 (O/S)

20

35%

7 (8)

No Burrow

None

N. americanus

1

30

50/50 (O/S)

20

30%

7 (8)

NA24

SVc

N. americanus

1

65

50/50 (O/S)

45

40%

7 (7)

NA28

V

N. americanus

5

30

50/50 (O/S)

20

35%

14 (21)

NA25 A-B

SV, H

N. americanus

5

65

50/50 (O/S)

43

37%

14 (14)

NA21 A-F

SV, SVb, SVc, Hc

N. americanus

1

30

50/50 (O/S)

20

35%

14 (14)

NA1

H

N. americanus

1

65

50/50 (O/S)

45

35%

14 (19)

No Burrow

None

Experiment 2: Sediment Composition

Species

Specimens

Tank Size

Sediment

Depth

% Moisture

Duration

Burrow #s

Burrow Architectures

F. penneri

5

30

50/50 (Sa/S)

16

50%

14 (94)

FP6 A-E

SV, OC (T1)

F. penneri

5

65

100 (S)

39

54%

7 (8)

FP7 A-H

V, Vh, SV,SVb, SVcb, J

F. penneri

4

30

50/50 (Sa/S)

19

70%

14 (54)

No Burrow

None

F. penneri

5

30

25/75 (Sa/S)

20

50%

14 (14)

FP5 A-G

V, SV, SVc, O (T1), O (T2), Ob (T2)

F. penneri

1

30

50/50 (Sa/S)

19

58%

14 (19)

FP4 A-B

V, H

F. penneri

1

30

25/75 (Sa/S)

20

52%

14 (41)

No Burrow

None

N. americanus

5

30

25/75 (Sa/S)

20

51%

14 (23)

NA32

SV

N. americanus

1

30

25/75 (Sa/S)

20

60%

14 (14)

NA31 A-B

O (T2), Om (T2)

N. americanus

1

65

100 (S)

39

60%

14 (51)

NA33 A-C

V, H

Experiment 3: Sediment Moisture

Species

Specimens

Tank Size

Sediment

Depth

% Moisture

Duration

Burrow #s

Burrow Architectures

F. penneri

5

65

50/50 (O/S)

45

49%

14 (63)

FP2 A-M

V, SV, J, Jb

F. penneri

1

65

50/50 (O/S)

45

50%

14 (13)

FP3 A-I

V, SV

F. penneri

1

65

50/50 (O/S)

45

50%

14 (14)

FP1

Hb

N. americanus

5

65

50/50 (O/S)

48

50%

14 (22)

NA29 A-C

V, H, O (T1)

N. americanus

1

65

50/50 (O/S)

50

50%

14 (21)

NA30 A-F

V, Vh, SV, SVc

 

TABLE 2. Quantitative measurements of burrow casts produced by Narceus americanus.

 
 

NA32

NA23

NA25A

NA21A

NA21B

NA21C

NA30B

NA30D

NA21D

NA24

NA21E

NA26A

NA30E

Architecture

SV

SV

SV

SV

SV

SV

SV

SV

SVb

SVc

SVc

SVc

SVc

                           

Surface Openings

1

1

1

1

1

1

1

1

1

1

1

1

1

                           

Maximum Depth (cm)

4.1

6.7

3.0

1.9

4.0

7.1

2.6

5.9

11.9

4.0

13.4

5.8

6.6

                           

Total Length (cm)

4.3

8.2

9.4

2.3

4.3

8.0

3.1

6.8

13.8

7.2

15.7

6.8

7.5

                           

Maximum Width (mm)

9.3

16.9

11.7

13.0

14.5

11.5

9.6

10.8

11.7

17.6

16.8

26.2

16.5

Minimum Width (mm)

8.5

8.0

7.8

10.5

11.0

8.8

9.5

7.5

8.6

7.1

7.2

11.2

6.0

Average Width (mm)

8.8

10.4

10.2

11.8

12.4

9.3

9.6

9.3

10.2

10.9

10.5

15.6

11.4

                           

Maximum Height (mm)

9.8

9.5

10.1

12.5

12.3

8.8

9.8

10.9

11.6

9.7

10.4

16.4

10.0

Minimum Height (mm)

8.6

7.3

5.7

11.8

10.5

7.4

8.2

8.1

6.3

6.0

5.8

8.9

7.0

Average Height (mm)

9.2

8.3

7.9

12.2

11.4

8.1

9.0

9.4

9.3

8.0

9.1

11.6

8.4

                           

Average W/H Ratio

1.0

1.3

1.3

1.0

1.1

1.1

1.1

1.0

1.1

1.3

1.2

1.4

1.3

                           

Maximum Circumference (cm)

3

4.1

3.7

4.2

4.3

3.1

2.9

3.5

3.6

4.6

4.3

8.4

4.7

Minimum Circumference (cm)

2.7

2.7

2.5

3.5

3.3

2.2

2.8

2.7

2.6

2.5

2.3

3.3

2.5

Average Circumference (cm)

2.8

3.1

3.1

3.9

3.7

2.7

2.9

3.1

3.1

3.3

3.2

4.9

3.5

                           

Maximum Slope (Degrees)

70

65

45

33

73

90

70

80

90

67

81

55

90

Minimum Slope (Degrees)

70

50

25

33

70

25

70

60

40

20

55

55

65

Average Slope (Degrees)

70

38

35

33

72

57

70

70

63

44

64

55

78

                           

Branching Angle (Degrees)

n/a

n/a

n/a

n/a

n/a

n/a

n/a

n/a

60

n/a

n/a

n/a

n/a

                           

Complexity

2.0

2.0

2.0

2.0

2.0

2.0

2.0

2.0

4.0

3.0

4.0

3.0

4.0

                           

Tortuosity

1.0

1.2

1.7

1.0

1.1

1.1

1.2

1.2

1.1

1.5

1.1

1.1

1.1

TABLE 2 (continued).

 
 

NA28

NA29C

NA30A

NA30C

NA33A

NA33B

NA30F

 

NA1

NA25B

NA33C

NA29B

NA21F

Architecture

V

V

V

V

V

V

Vh

 

H

H

H

Hb

Hc

                           

Surface Openings

1

1

1

1

1

1

1

 

1

1

1

1

1

                           

Maximum Depth (cm)

2.4

1.9

2.5

2.3

3.1

2.2

5.9

 

2.8

2.5

3.0

3.4

2.9

                           

Total Length (cm)

3.0

1.9

2.5

2.6

3.1

3.0

11.8

 

7.0

8.4

6.8

10.7

10.0

                           

Maximum Width (mm)

14.0

11.7

9.3

10.4

12.0

13.8

10.4

 

10.2

11.0

11.8

10.6

16.5

Minimum Width (mm)

14.0

11.7

7.0

9.3

10.4

13.0

7.2

 

7.4

9.0

7.0

7.4

6.8

Average Width (mm)

14.0

11.7

8.2

9.9

11.2

13.4

9.1

 

8.9

10.1

9.6

8.4

11.6

                           

Maximum Height (mm)

13.8

9.0

8.8

9.3

14.7

14.1

9.6

 

9.8

9.1

11.7

9.2

11.5

Minimum Height (mm)

9.5

9.0

7.7

8.0

11.1

10.6

6.7

 

6.3

5.8

8.5

6.1

8.0

Average Height (mm)

11.7

9.0

8.3

8.7

12.9

12.4

8.3

 

8.3

8.0

9.8

7.8

9.7

                           

Average W/H Ratio

1.2

1.3

1.0

1.1

0.9

1.1

1.1

 

1.1

1.3

1.0

1.1

1.2

                           

Maximum Circumference (cm)

5

3.5

2.8

3.1

4.3

4.4

3.2

 

3.2

3.3

4.3

3

4.3

Minimum Circumference (cm)

3.7

3.5

2.4

2.8

3.5

3.9

2.1

 

2.2

3

2.3

2.5

2.3

Average Circumference (cm)

4.4

3.5

2.6

3.0

3.9

4.2

2.8

 

2.8

3.2

3.1

2.7

3.4

                           

Maximum Slope (Degrees)

85

80

90

80

90

90

90

 

60

50

80

60

55

Minimum Slope (Degrees)

85

80

90

80

90

90

0

 

15

0

50

60

0

Average Slope (Degrees)

85

80

90

80

90

90

45

 

45

25

65

75

25

                           

Branching Angle (Degrees)

n/a

n/a

n/a

n/a

n/a

n/a

n/a

 

n/a

n/a

n/a

90

n/a

                           

Complexity

2.0

2.0

2.0

2.0

2.0

2.0

2.0

 

2.0

2.0

2.0

4.0

4.0

                           

Tortuosity

1.3

1.0

1.0

1.1

1.0

1.0

2.0

 

3.5

5.6

2.3

1.5

1.2

TABLE 2 (continued).

 
 

NA29A

NA26B

NA31A

NA31B

 

Total Mean

Architecture

O (T1)

O (T1)

O (T2)

O (T2)

   
             

Surface Openings

1

1

2

1

 

1

             

Maximum Depth (cm)

3.4

6.9

2.5

1.9

 

4.4

             

Total Length (cm)

10.6

12.5

12.3

10.0

 

7.4

             

Maximum Width (mm)

12.9

18.0

13.4

10.9

 

13.2

Minimum Width (mm)

8.3

7.4

8.7

9.1

 

8.8

Average Width (mm)

10.5

10.9

10.4

10.1

 

10.6

             

Maximum Height (mm)

9.6

11.3

12.7

11.7

 

11.0

Minimum Height (mm)

6.0

6.7

9.3

8.9

 

7.9

Average Height (mm)

8.2

9.3

10.4

9.9

 

9.5

             

Average W/H Ratio

1.3

1.2

1.0

1.0

 

1.1

             

Maximum Circumference (cm)

3.5

4.8

3.9

3.6

 

4.0

Minimum Circumference (cm)

2.7

2.3

2.9

3.1

 

2.8

Average Circumference (cm)

3.1

3.4

3.5

3.3

 

3.3

             

Maximum Slope (Degrees)

60

60

90

20

 

71

Minimum Slope (Degrees)

15

60

0

20

 

47

Average Slope (Degrees)

38

60

43

20

 

59

             

Branching Angle (Degrees)

n/a

n/a

n/a

n/a

 

75

             

Complexity

3.0

3.0

6.0

2.0

 

2.6

             

Tortuosity

43.0

39.0

1.8

99.0

 

7.6

TABLE 3. Quantitative measurements of burrow casts produced by Floridobolus penneri.

 
 

FP2C

FP3C

FP3D

FP3E

FP3F

FP3G

FP3H

FP3I

FP2E

FP2F

FP2G

FP6A

Architecture

SV

SV

SV

SV

SV

SV

SV

SV

SV

SV

SV

SV

                         

Surface Openings

1

1

1

1

1

1

1

1

1

1

1

1

                         

Maximum Depth (cm)

3.5

1.7

3.3

3.3

3.6

6.8

6.5

8.2

7.9

6.9

4.4

1.8

                         

Total Length (cm)

4.0

1.7

4.2

3.3

4.2

7.4

6.8

8.6

9.6

9.5

10.0

4.3

                         

Maximum Width (mm)

13.1

9.4

14.0

14.1

14.2

10.8

12.8

12.8

12.5

12.2

6.5

17.0

Minimum Width (mm)

11.0

9.4

9.6

13.2

12.4

8.8

10.2

10.3

9.2

9.7

0.9

10.0

Average Width (mm)

12.1

9.4

11.1

13.7

13.2

10.1

11.4

11.4

11.0

10.8

4.0

13.7

                         

Maximum Height (mm)

14.3

10.6

11.6

12.0

11.1

12.0

12.4

12.5

13.3

11.4

7.8

10.7

Minimum Height (mm)

10.0

10.6

8.6

8.3

9.2

8.7

10.8

8.5

8.1

8.7

3.4

10.0

Average Height (mm)

12.8

10.6

10.0

10.2

10.4

10.5

11.5

10.5

10.9

10.1

5.8

10.3

                         

Average W/H Ratio

1.0

0.9

1.1

1.4

1.3

1.0

1.0

1.1

1.0

1.1

0.7

1.3

                         

Maximum Circumference (cm)

4.4

3.2

4

4.4

4

3.7

4

4.1

3.9

3.7

3.7

4.5

Minimum Circumference (cm)

3.4

3.2

3.1

3.7

3.8

2.9

3.1

3.2

3.2

2.9

2.4

2.8

Average Circumference (cm)

4.0

3.2

3.4

4.1

3.9

3.3

3.6

3.6

3.6

3.4

3.2

3.7

                         

Maximum Slope (Degrees)

65

75

68

65

75

78

70

65

65

55

78

20

Minimum Slope (Degrees)

65

75

68

65

75

78

70

65

65

55

78

20

Average Slope (Degrees)

65

75

68

65

75

78

70

65

65

55

78

20

                         

Branching Angle (Degrees)

n/a

n/a

n/a

n/a

n/a

n/a

n/a

n/a

n/a

n/a

n/a

n/a

                         

Complexity

2.0

2.0

2.0

2.0

2.0

2.0

2.0

2.0

2.0

2.0

4.0

2.0

                         

Tortuosity

1.0

1.0

1.2

1.0

1.2

1.1

1.0

1.1

1.1

1.2

1.1

1.4

TABLE 3 (continued).

 
 

FP3A

FP5B

FP5C

FP5D

FP7A

FP7C

FP7E

FP7F

FP7H

FP5E

Architecture

SV

SV

SV

SV

SV

SV

SV

SVb

SVcb

SVc

                     

Surface Openings

1

1

1

1

1

1

1

1

1

1

                     

Maximum Depth (cm)

1.5

2.8

4.2

5.7

2.1

4.1

8.7

8.0

7.4

5.2

                     

Total Length (cm)

3.3

3.4

4.9

7.7

2.4

5.5

9.2

12.4

18.6

6.8

                     

Maximum Width (mm)

8.2

15.9

12.9

16.3

11.3

16.0

15.0

18.3

30.1

16.4

Minimum Width (mm)

7.3

14.5

8.0

12.7

10.3

9.1

9.0

10.1

8.7

10.4

Average Width (mm)

7.8

15.2

11.0

13.9

10.8

12.8

12.3

13.6

16.8

12.7

                     

Maximum Height (mm)

8.5

19.6

11.8

16.3

12.0

14.0

na

12.8

19.0

24.7

Minimum Height (mm)

7.6

13.3

10.9

11.8

10.8

10.7

na

10.3

8.7

11.5

Average Height (mm)

8.1

16.5

11.5

13.9

11.4

12.7

10.0

11.3

13.4

15.6

                     

Average W/H Ratio

1.0

1.0

1.0

1.0

0.9

0.9

1.2

1.2

1.3

0.9

                     

Maximum Circumference (cm)

2.7

5.7

4.1

5.2

3.6

4.5

3.9

4.5

8.1

6.2

Minimum Circumference (cm)

2.6

4.6

3.2

3.3

3.1

3

2.9

3.1

2.6

3.3

Average Circumference (cm)

2.7

5.2

3.7

4.5

3.4

3.9

3.6

4.0

5.0

4.7

                     

Maximum Slope (Degrees)

45

45

70

55

75

75

90

90

90

65

Minimum Slope (Degrees)

45

45

70

55

75

50

65

50

20

45

Average Slope (Degrees)

45

45

70

55

75

62

78

70

50

55

                     

Branching Angle (Degrees)

n/a

n/a

n/a

n/a

n/a

n/a

n/a

20

55

n/a

                     

Complexity

2.0

2.0

2.0

2.0

2.0

2.0

2.0

3.0

4.0

3.0

                     

Tortuosity

1.6

1.1

1.1

1.2

1.1

1.3

1.1

1.1

1.5

1.3

TABLE 3 (continued).

 
 

FP3B

FP2A

FP2B

FP2D

FP4A

FP5A

FP7B

FP2M

FP7D

Architecture

V

V

V

V

V

V

V

V

Vh

                   

Surface Openings

1

1

1

1

1

1

1

1

1

                   

Maximum Depth (cm)

2.2

4.1

3.1

6.0

3.3

2.1

3.6

3.6

6.9

                   

Total Length (cm)

2.2

4.1

3.1

6.7

3.6

2.1

4.7

3.6

8.4

                   

Maximum Width (mm)

12.6

10.6

15.0

15.6

16.5

16.9

14.5

23.3

14.4

Minimum Width (mm)

9.8

9.2

13.6

8.6

12.3

10.3

7.6

12.1

10.3

Average Width (mm)

11.2

9.9

14.3

13.2

14.7

13.6

12.0

16.6

12.1

                   

Maximum Height (mm)

9.7

11.9

15.8

13.3

13.4

15.4

13.7

23.0

11.3

Minimum Height (mm)

9.1

10.0

14.0

9.6

10.5

12.0

11.6

12.7

9.8

Average Height (mm)

9.4

11.1

14.9

11.5

12.2

13.7

12.7

16.7

11.0

                   

Average W/H Ratio

1.2

0.9

1.0

1.1

1.2

1.0

0.7

1.0

1.1

                   

Maximum Circumference (cm)

3.6

3.8

5.2

5.6

5.5

4.6

4

6.5

3.8

Minimum Circumference (cm)

3

3.4

4.6

2.6

3.5

3.4

3.2

3.8

3.3

Average Circumference (cm)

3.3

3.6

4.9

4.4

4.6

4.0

3.6

5.2

3.6

                   

Maximum Slope (Degrees)

90

85

90

85

90

90

90

90

90

Minimum Slope (Degrees)

90

85

90

60

90

90

40

90

55

Average Slope (Degrees)

90

85

90

73

90

90

65

90

73

                   

Branching Angle (Degrees)

n/a

n/a

n/a

n/a

n/a

n/a

n/a

n/a

n/a

                   

Complexity

2.0

2.0

2.0

2.0

2.0

2.0

2.0

2.0

2.0

                   

Tortuosity

1.0

1.0

1.0

1.1

1.1

1.0

1.2

1.0

1.2

TABLE 3 (continued).

 
 

FP2I

FP2J

FP2K

FP2L

FP7G

FP2H

 

FP4B

FP1

 

FP5F

FP6B

FP5G

 

Total Mean

Architecture

J

J

J

J

J

Jb

 

H

Hb

 

Ob (T1)

Oc (T1)

Ob (T2)

 
                               

Surface Openings

1

1

1

1

1

1

 

1

1

 

1

1

1

 

1

                               

Maximum Depth (cm)

6.5

7.8

8.1

2.9

8.4

5.8

 

3.1

2.8

 

7.7

4.4

1.1

 

4.8

                               

Total Length (cm)

10.7

11.9

11.8

7.5

14.7

13.0

 

11.1

11.1

 

17.5

13.0

10.1

 

7.6

                               

Maximum Width (mm)

11.5

16.6

12.4

11.4

13.0

13.0

 

13.6

13.5

 

12.5

24.8

11.9

 

14.4

Minimum Width (mm)

8.4

9.3

9.5

7.9

9.0

7.2

 

10.0

7.5

 

9.1

10.0

9.0

 

9.7

Average Width (mm)

9.9

12.0

11.0

9.4

11.3

10.8

 

11.6

9.8

 

10.8

17.7

10.2

 

11.9

                               

Maximum Height (mm)

13.0

16.5

14.8

12.1

15.7

14.7

 

11.4

7.0

 

13.7

16.0

11.2

 

13.5

Minimum Height (mm)

9.2

9.2

10.3

7.4

10.4

7.7

 

7.5

5.1

 

10.0

7.7

6.2

 

9.5

Average Height (mm)

10.7

11.8

13.0

9.5

12.4

11.0

 

10.0

6.1

 

12.2

12.1

8.3

 

11.4

                               

Average W/H Ratio

0.9

1.0

0.9

1.0

0.9

1.0

 

1.2

1.6

 

0.9

1.5

1.3

 

1.1

                               

Maximum Circumference (cm)

3.7

5.3

4.2

3.5

4.2

6.4

 

4.3

3.4

 

4.1

6.1

3.5

 

4.5

Minimum Circumference (cm)

2.9

2.8

3

2.6

3.1

2.4

 

3.1

2.2

 

3.2

2.6

2.7

 

3.1

Average Circumference (cm)

3.2

3.8

3.8

3.1

3.9

3.7

 

3.6

2.7

 

3.7

4.4

3.1

 

3.8

                               

Maximum Slope (Degrees)

80

90

55

55

75

65

 

70

50

 

85

90

20

 

72

Minimum Slope (Degrees)

15

50

50

45

75

25

 

0

25

 

85

40

20

 

58

Average Slope (Degrees)

47

70

53

50

75

45

 

35

38

 

85

65

20

 

65

                               

Branching Angle (Degrees)

n/a

n/a

n/a

n/a

n/a

70

 

n/a

10,45

 

12, 25

n/a

10

 

31

                               

Complexity

2.0

2.0

2.0

2.0

2.0

3.0

 

2.0

6.0

 

6.0

4.0

4.0

 

2.5

                               

Tortuosity

2.2

2.0

2.0

5.8

2.4

1.4

 

4.3

1.1

 

17.6

53.0

1.4

 

3.1

 

 

TABLE 4. Rs values of the Spearman's rank correlation tests between sediment clay and moisture content and quantitative burrow properties. The number of burrow openings rarely varied between burrow casts and were, therefore, excluded from the analysis.

 

Narceus americanus

 

Length

Depth

Width

Height

Circum.

Slope

Complexity

Tortuosity

Clay

-0.31

-0.10

0.01

0.42

-0.13

-0.39

-0.28

-0.30

Moisture

-0.22

-0.33

-0.13

0.41

0.04

0.46

-0.10

-0.11

                 

Floridobolus penneri

 

Length

Depth

Width

Height

Circum.

Slope

Complexity

Tortuosity

Clay

0.15

0.19

0.24

0.36

0.22

0.03

-0.17

0.14

Moisture

0.03

-0.07

0.26

-0.01

0.04

-0.01

0.09

0.11

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