9. Morphometric Adaptations of the Mangrove
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9. MORPHOMETRIC ADAPTATIONS OF THE MANGROVE LOBSTER THALASSINA ANOMALA HERBST (DECAPODA, THALASSINIDAE) TO A BURROWING LIFE-STYLE Gunasagaran Pillai School of Pure and Applied Sciences The University of the South Pacific P.O. Box 1168, Suva, FIJI ABSTRACT Morphometric relationships of the carapace, abdomen and the first pleopods of the mangrove lobster Thalassina anomala have been examined . Results indicate that these may be satisfactorily described by the simple linear equation, y = a + bx. The functional morphology of these structures and their relative growths suggest that the animal is admirably adapted to a burrowing life style. INTRODUCTION Studies of relative growth by morphometric analysis in Crustacea have been largely confined to the Brachyura (Teissier, 1960; Hartnoll, 1978,1982) With the possible exception of f.!!.lll.!!..!!..!!.~~.!!. australiensis (Hailstone & Stephenson, 1965), such analyses have not been extended to include the Anomura. In the course of growth it is common for some dimensions to increase at different rates from others, as a result of which there is change in proportion with size. This is called relative growth. Crustacea were one of the earliest groups to lend themselves to measurement of relative growth because of their hard integument, sexual dimorphism and variance in appendicular dimensions. During the early stages in the study of relative growth Huxley (1924) demonstrated that nearly all examples could be satisfactorily described in accordance with the simple allometricy equation y = Axo, where x is the reference S. Pac. J. Nat. Sci., 1987, Vol. 9 9-20 10. dimension, y the variable dimension, A the y- intercept, and B is the regression coefficient. This equation implies that the two dimensions grow relative to each other at a constant rate, defined by the parameter B, and it is known as the relative growth rate or the level of allometry (Hartnoll, 1982). If'" exceeds 1, growth is positively allometric, isometric when '" 1, and negatively allometric when", <1. Expressed logarithmically the equation becomes: log y = log A + '" log x. If log y is plotted as a function of log x, a straight line is obtained, the slope of which has the value "'. In this paper morphometric relationships of the carapace, abdomen and first pleopods of the mangrove lobster Thalassina anomala Herbst. are discussed in relation to its burrowing life-style. This animal inhabits the littoral fringe of estuarine shores of the tropics where it is usually found in association with mangrove swamps. In the clayey-silty deposits of the swamp, it leads a cryptic, burrowing mode of life, occasionally.coming to the surface to unload pallets of mud. The manner in which the underground sediment is brought outside, with the aid of its first two pairs of pereiopods and the third maxi llipeds, is analogous to the actions of the scoop of a mechanical front-end loader. The tunnel is usually tortuous and the shaft tends to become vertical as it reaches the entrance. Such a way of life requires certain specialized morphological adaptations. MATERIALS AND METHODS All specimens of I. anomala were collected from the environs of the o 0 Rewa delta area (1807'S, 178 31'E), Fiji Islands. Larger specimens were caught by means of traditional traps (Pillai, 1985), and juveniles were dug out with a spade. Despite laboriously digging up more than a hundred mounds, only five juveniles (none of the. females) were obtained. As a result of this, the samples did not include a sufficient number of smaller individuals. Measurements were taken to the nearest millimetre with the aid of a divider and a 11. ruler. Linear plots of carapace length-width, carapace length-abdomen length, abdomen length-abdomen width, and abdomen length-first pleopod len gth were made. In all cases the correlation was significant (P < 0.01). Differences in morphometric relationships based on the linear regressions between the sexes were tested by analysis of covariance (ANOCOVA). The level of allometry was determined by testing the regression coefficient or slope (a) of the log-transformed regressions (see Hartnoll, 1982) against the isometric slope of 1 with Student's test. RESULTS AND DISCUSSION A. RELATIONSHIP BETWEEN CARAPACE LENGTH AND WIDTH As there was no evidence of sexual dimorphism with respect to carapace dimensions, the carapace lengths and widths for a range of size of both sexes were pooled for analysis. From these data, carapace width was plotted as a function of carapace length (Fig . 1). The slope a of the log-transformed regression was found to be 1.00 (Table I), indicating that carapace growth is isometric. Isometric growth is functionally significant because a cha~ge in shape (for example, a widening of the carapace) would have deleterious effects on the burrowing activities of the mangrove lobster. The animal, being a front-burrower, moves only forwards or backwards through the burrow. A positively or a negatively allometric growth, i.e., a >1, or a <I, respectively, could alter the basically cylindrical cross-sectional nature of the body of the burrower. This presumably would necessitate the widening of the tunnel which would be energetically wastef ul. In brachyurous side- burrowers (e.g. Macrophthalmus) on the other hand, there is a change in carapace shape during growth; it becomes shorter and wider (Barnes, 1968). This is adaptive, for an increase in width does not alter the inherent cylindrical shape of this side-burrower. 12. o o o 4 o o o - -0.06 + (0527 ± 0 . 045)x 0.99 1 14 o o o o 2 4 CA RAPA CE LEN GTH ( CM) Figure 1. Relationship between carapace length and width in I. anoma1a. C'"l JJEBi1S * JJEBi1S ** IIEBIIS ...... Table 1. Regressicn analysis of nDrJiuJetdc data of I: anarala (ns not significant (p)O.05) ; p<O .05; p<O .Ol). Linear Regressicn Log-transforne:l ANOCOVA (y = 8 + bx) Regressiro (y = A+X) Slope J 8 b r A B r Carp.ris:n of variances Carp.ris:n of slopes Ccnp.rign of Elevaticn ~e length nale and femle -{l.a; 0.527 -IO.'1:l1 -0.663 1.CXl3 0.993 v pooled CaraJBCe width n = 14 CaraJBCe l ength nale v n = 23 -{l. ~ 1.836 -10. 994 0.417 l.ai9 O.~ F = 1.87 ns F = 0.48 ns F = l.01 ns (d.L = 34 ,22) (d.L = 1,54) (d .L = l,ssJ abdrnen l ength femle -1.11 1.918 -IO .9B4 0.391 um 0 .993 n = 35 Abdrnen length nale v n = 22 0.05 0.197 -IO. <m -1.4B7 0.951 O.<m F = 6 .73** F = 11.91 ** F = 23** abdaren width (d.f. = 21, 27) (d.f. = I , IKJ) (d.L = 1,4/ femle -{l.35 0.271 -10.% -1.853 1. 1~ 0.992 n = 2B Abdrnen l ength nale -{l.a; 0.151 0.994 -2.122 1.077 0.997 ... v n = 14 F = 4.67** F = 4.'ll** F = 683.37 first plropod (d.f . = 16, 13) (d.L = 1, 127) (d.f. = 1,2E length femle -{l.B2 0 .115 -10 .761 -4.713 I. ~ 0. /89 n = 17 14. 18 n = 0.69 + ( 1.836 ± 0.042)x 16 ,JI r = 0.994 n = 23 14 12 ~ 1 y = -1.11 + ( 1. 918 ± 0.1 24lx 0.98 4 n = 35 4 2 4 10 12 CA RAPAC E LE NG TH (CM) Figure 2. Relationship between carapace len g th and abdomen length in I. anomala. 15. B. RELATIONSHIP BETWEEN CARAPACE LENGTH AND ABDOMEN LENGTH Abdomen length was plotted against carapace length for the two sexes separately (Fig. 2). The value of {l (of the log-transformed regression) was found to be equal to 1.09 for females, and 1.07 for males. These values did not indicate significant positive allometry (P>O.Ol). ANOCOVA (Table 1) showed that there was little difference in the magnitudes of the two variables, i.e. carapace length and abdomen length, between sexes. Hence, total lengths of larger individuals of either sex are about the same. A lack of dimorphism in size between them may be adaptive for it might enable both sexes to occupy the same burrow. C. RELATIONSHIP BETWEEN ABDOMINAL LENGTH AND WIDTH A linear plot of abdomen width against abdomen length is shown in Fig. 3. Log-transformed regression gave a values of 1.17 and 0.95 for females and males, respectively (Table 1). Hence, while abdomen growth is positively allometric for females, it is slightly negatively allometric for males, the sexual difference being significant. The shape of the a bdomen in the mangrove lobster is basically the same in both sexes, although its relative width varies somewhat between sexes. The abdominal length-width ratios of the males and females were found to be approximately 1:5 and 1 :4, respectively. The female abdomen is slightly wider than that of the male. The abdomen of brachyurans, on the other hand, shows striking dimorphism. Wider abdomen of female decapods is of functional significance since the developing eggs are carried attached to the pleopods after extrusion. The elongate abdomen of the mangrove lobster, which constitutes 65% of the total length (unpublished observation), allows a large number of eggs to be borne (Pillai, 1982) without affecting the burrowing activity of the female. A significant increase in a-~ d" y = 0.05 + (0. 197 0.014lx r = 0.988 n = 22 ~ J /"" ~. i: 0 '"z ~ 2 0 0 <n "" I • ~ 0.098)x 0. 986 • 28 I 4 10 12 14 16 18 ABDOMEN LENGTH (eM) Figure 3. Relationship between abdomen length and width in I. anomala. 17. abdomen width would have an adverse effect on the burrowing ability of the animal, i.e., the burrows would have to be widened.