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Allometric Growth of the Freshwater Crab Potamon algeriense (Bott, 1967) (Decapoda, Brachyura, Potamidae) in Oued Zegzel, a Mountain Stream, in the Northeast of Morocco

Allometric Growth of the Freshwater Crab Potamon algeriense (Bott, 1967) (Decapoda, Brachyura,... Hindawi International Journal of Zoology Volume 2019, Article ID 5168639, 8 pages https://doi.org/10.1155/2019/5168639 Research Article Allometric Growth of the Freshwater Crab Potamon algeriense (Bott, 1967) (Decapoda, Brachyura, Potamidae) in Oued Zegzel, a Mountain Stream, in the Northeast of Morocco 1 1 2 1 Soufiane Fadlaoui , Mohammed Mahjoub, Ouahid El Asri, and Mohammed Melhaoui Mohamed First University, Faculty of Sciences, Department of Biology, Laboratory of Water, Environment, and Sustainable Development, P.B. 717, Oujda, Morocco Biochemistry and Biotechnology Laboratory, Mohamed First University, Oujda, Morocco Correspondence should be addressed to Soufiane Fadlaoui; soufiane.fadlaoui@gmail.com Received 30 January 2019; Revised 4 April 2019; Accepted 27 June 2019; Published 1 August 2019 Academic Editor: o Th mas Iliffe Copyright © 2019 Soufiane Fadlaoui et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. A morphometric study has been carried out for the first time on the population of the freshwater crab Potamon algeriense inhabiting Oued Zegzel, a tributary of the Lower Moulouya River, in the northeast of Morocco. Crabs were collected monthly over one year (October 2017 to September 2018) by excavating burrows and searching under blocks. A total of 669 crabs were obtained, 291 females and 378 males. Regression analysis was performed among carapace width (CW), as the reference dimension, and wet weight (WW),carapace length (CL),lengthand widthofabdomen (ALandAW), andlengthand widthofthecheliped (ChL and ChW) were chosen as dependent variables, using the allometric method. Based on Somerton’s technique, the onset of sexual maturity wasestimatedto occurataround27mmCWfor malesand32mmCW forfemales.Thegrowthpatternsrecordedfor P. algeriense are associated with the species reproductive strategy, i.e., preparation of body parts involved in female acquisition and egg incubation, like the male cheliped and the female abdomen, respectively. Among the 291 females sampled, 10.20% were left-handed and 89.80% were right-handed, while among the 378 males examined, 10.32% were left-handed and 89.68% were right-handed. It was concluded that the cheliped width and the abdominal width of P. algeriense are the morphometric variables most appropriate to estimate the size at the beginning of the sexual maturity for males and females of this species, respectively. 1. Introduction and to establish sexual maturity as well as contributing to the identification of “handedness” occurrence [9–12]. Allometry, which is the oldest of the approaches and still Size at which sexual maturity is attained is important widely applied in biology, is a well-known phenomenon in crab life history, because both age and size at maturity concerning the study of the relationship between the size and are associated with a species’ reproductive output [13]. An function of organs of the body and growth or size of the whole assemblage of morphological, physiological, and behavioral body [1]. Allometry is oeft n divided into three principal types: changes through which immature individuals become able ontogenetic (when shape changes with ontogenetic stage or to produce and transfer gametes marks the onset of maturity age), static (when shape correlates with size independently [14]. of age), and evolutionary (when shape correlates with size eTh growth pattern of some specicfi body parts such as among species) [2]. chelipeds, abdomen, and gonopods shows variations in the Relative growth in brachyuran crabs has been commonly degree of allometry during the course of development, which studied since the earlier decades of the century [3–5], but only may coincide with gonad maturation, providing an important in more recent work were these allometric patterns reviewed estimate for the size at which these animals are ready for for the group [6]. It can be used in species identification [7, 8] reproduction [15, 16]. O. Bouizi Oued Cherraa 2 International Journal of Zoology Nador Mediterranean Sea Plain of Triffa W E W E Berkane Midar S S Oujda O. Bou Rdim Aknoul Oued Moulouya Taourirt O. Ouizert O. Charef Taza km O. Charef 0 2 4 Ifrane O. Azreg Azrou Missour Khenifra Midelt Figure 1: Localization of the study area, Oued Zegzel, Morocco. Crabs of the infraorder Brachyura are among the most stream, in the northeast of Morocco, focusing on the mor- diverse groups of crustaceans with more than 7000 described phometric relationships between carapace width and the size species in 98 families, occurring in various habitats: marine, of other body components. Through the analyses of these freshwater, and terrestrial [17–20]. parameters, it is possible to characterize the allometry levels Freshwater crabs are found exclusively in freshwater or and to make inferences about the sizes at which males and terrestrial ecosystems and never enter brackish or marine females reach sexual maturity, thus evidencing their puberty waters for reproduction [21]. Approximatively, 1400 species molt. The frequency of handedness (major cheliped) in each are known with additional species being regularly discovered sex and its growth are also studied. reviewed in [16, 17]. Potamon algeriense belongs to the family of the Potamidae 2. Material and Methods which is the largest of all freshwater crab families and com- 2.1. Study Area. The study was performed in Oued Zegzel prises 95 genera and over 505 species distributed through- ∘ 󸀠 ∘ 󸀠 (34 50 29.4 N, 2 21 19.8 W), a tributary of the Lower out the southern Palaearctic and Oriental zoogeographical Moulouya River, in the northeast of Morocco (Figure 1). eTh regions from Morocco as far east as Japan and as far south region is characterized by a Mediterranean climate. as Indonesia [21, 22]. In fact, it is the only Maghrebian Shrublands that mark the study area were composed representative of this family. eTh species could be found mainly of Nerium oleander, Tamarix africana, Salix pedi- exclusively in three countries: Morocco, Algeria, and Tunisia cillata, Rubus ulmifolius, Rubus ulmifolius,and Crataegus [23]. In Morocco, the species has been reported from the monogyna.Thus,agroup ofsubmerged plants ( Nasturtium north in the watershed of the Oued Laou near Chefchaouen, officinale , Veronica catenata,and Scrophularia aquatica)is from the northeast in watershed of Moulouya, and from individualized on the banks of the Oued, while in the central the Middle Atlas in the Oued Oum er Rbia watershed near waters Potamogeton natans or Potamogeton pectinatus are Khenifra. oen ft overgrown. Despite the wide distribution of P. algeriense,itsoccur- rence drastically declined during the last two decades due to human impacts (sewage, industrial waste, and agricultural 2.2. Crab Sampling and Measurement. The crabs were col- pesticides). Consequently, P. algeriense is included in the lected monthly over one year (October 2017 to September IUCN red list as least concern species. Comprehensive 2018)byexcavatingburrows andsearchingunderblocks.The biological information about this species is incomplete, as crabs could be caught by plunging an arm into the burrow. previous studies have only treated its geographic distribution However, as the depth of the burrow can reach more than 50 and taxonomic status [24]. cm it was not always possible to catch the crabs before they Thisstudy aims toanalyzeforthefirsttimetherelative escaped to the greater depths. Consequently, the individuals growth of P. algeriense, inhabiting Oued Zegzel, a mountain assessed here represent a random sample. O. Melloulou Atlantic Ocean O. Moulouya Oued Zegzel O. Ouizert Oued M. Idriss 󸀠󸀠 󸀠󸀠 International Journal of Zoology 3 ChL Female Male C CW W A A A AW W CL AL C ChW hW AL AW Figure 2: Position of measurements in Potamon algeriense. CW: carapace width; CL: carapace length; ChL: cheliped length; ChW: cheliped width; AW: abdomen width; AL: abdomen length. All measurements were made to the nearest (0.01 mm) 3. Results using a vernier caliper: carapace width (CW) used as the A total of 669 specimens of P. algeriense were collected, and reference dimension, measured at the widest part; carapace their CW sizes ranged from 5.40 to 26.9 mm for juvenile length (CL); abdominal width (AW); abdominal length (AL) males (N=323), from 27 to 60.30 mm CW for adult males (taken from the widest segment); and cheliped width (ChW) (N=55), 11.80 to 31.9 mm for juvenile females (N=158), and and length (ChL). eTh position of these measurements is 32 to 50.10 mm for adult females (N=133). indicated in Figure 2. According to Somerton’s technique [28, 29], the rela- The specimens were also weighted, wet weight (WW), tionships that best indicate the change in the allometric using an analytical pocket balance to the nearest 0.01 g. coefficient between juveniles and adults phase and the dis- Sex determination was based on the morphology of the tinction between male and female growth are CW vs. AW abdomen (considerably wider in females than in males) and for females and CW vs. ChW for males. In fact, the size the number of pleopods. The presence of eggs and juvenile at which 50% of the females reached maturity is 32 mm in the abdomen of females was also recorded. Chelipeds were of CW based on the CW vs. AW relationship. At this size, designated as major or minor by their size. females present a noticeable growth of the abdomen width Only complete and sexed specimens were used in this resultinginanincreaseoftheallometric coecffi ient(1.38). study. On the other hand, the relationship between ChW and CW in males showed a remarkable increase of the cheliped width 2.3. Statistical Analyses. Relative growth was examined fol- soon aer ft the puberty molt, which may occur in crabs of lowing the method of Huxley [25] to describe most patterns approximately 27 mm CW, resulting in an increase of the of relative growth in animals in general and in brachyuran allometric coefficient (1.38) (Figure 3). crabs especially [15, 26, 27]. The regressions parameters (slope and intercept) and the The logarithmic transformation log Y = log a + b log X relative growth equations of the morphometric relationships was linearized from the traditional allometric growth equa- obtained for both sexes in juveniles and adults, and their tion Y = aX , where (Y) and (X) are morphological dimen- departures from isometry (t-test) are shown in Tables 1 and sions and (a) and (b) are growth constraints. eTh computer 2, respectively. programs Mature Iand II[28,29]wereusedtodelimit the er Th elationshipsofWW,AW,ChW,andChLwithCW size at which 50% of the individuals reached maturity. as an independent variable indicate that there are significant Analysis of covariance (ANCOVA) was performed to differences in growth between juveniles and adults in both compare slopes and intercepts of the obtained regressions sexes. within each allometric relationship and between sexes [30, Concerning handedness, right-handed considerably out- 31]. Carapace width (CW) was chosen as the reference numbered left-handed ones; from 669 crabs examined, dimension (independent variable), since this has been widely 10.31% (69 of 669) had the left propodus and dactylus used for recent studies on decapods including the Potamidae enlarged (left-handed) and 89.69% (600 of 669) had the [32]. right propodus and dactylus enlarged (right-handed). No Departures from isometry (Ho: b= 3) for wet weight and homochelous individuals were found. Among the 291 females (Ho: b= 1) for the other measurements were tested using a sampled, 10.20% (30 of 291) were left-handed and 89.80% Student’s t-test on the obtained slope values (𝛼=0.05) [31]. (261 of 291) were right-handed, while among the 378 males If b> 1, then the dimension (Y) increases in size relatively examined, 10.32% (39 of 378) were left-handed and 89.68% more rapidly than the reference dimension (X) does and (339 of 669) were right-handed. No significant difference in growth is said to be positively allometric. A condition of b< handedness between sexes was observed (𝜒 =4.16; P> 0.05). 1 indicates negative allometry, and b = 1 indicates a condition of isometry that means there is no change in the relative 4. Discussion shape with increasing size. Handedness between sexes was analyzed using a two-tailed𝜒 test to determine whether the In crustaceans, some aspects such as gonadal development, frequencies of right- and left-handed males were similar. presence of ovigerous females, and morphometric data are 4 International Journal of Zoology 1,60 Males Females 1,40 Adults 1,40 1,20 1,00 Adults 1,20 Juveniles ,80 Juveniles 1,00 ,60 ,40 ,80 ,20 ,60 ,00 1,00 1,20 1,40 1,60 ,75 1,00 1,25 1,50 1,75 Log Carapace Width (mm) Log Carapace Width (mm) (a) (b) Figure 3: Dispersion points and adjusted curve of the relationships: (a) ChW vs. CW; (b) AW vs. CW for males and females of Potamon algeriense, respectively. used to calculate the mean size at the onset of sexual maturity. growth of cheliped dimensions (generally cheliped width) to In fact, the estimation of puberty size may vary according characterize a possible sexual dimorphism and maturation in to the body part analyzed. In the present study, the mean crabs [6, 34]. size of maturity was calculated according to the relationships The size at the onset of sexual maturity of P. algeriense of ChW vs. CW and AW vs. CW for males and females, from Oued Zegzel was greater in females than in males, respectively. es Th e relationships provide the best tfi of sexual andsimilarresultshavebeenfound inthepopulations of maturity of P. algeriense.Gherardiand Micheli[33]remarked Mithraculus forceps, Mithrax tortugae, and Grapsus adscen- that freshwater crabs mate during the intermolt phase when sionis [40–42]. On the contrary, the onset of sexual maturity the male grabs the female within its chelipeds, pointing out for the populations of T. u fl viatilis and D. pagei [39, 43] from the importance of chelipeds for the reproductive process and south-eastern Brazil occurred at the same size interval, as confirming the choice of this body part as the reference for estimated based on chelipeds and abdomen growth for males the analysis of size at sexual maturity. and females, respectively. Delayed sexual maturity could The patterns of relative growth by abdomen, cheliped, and allow females of P. algeriense to live longer, grow larger, and heterochely in P. algeriense follow those already remarked in therefore have higher rate of fecundity [44]. Furthermore, the most crab species studied so far [6], and the adaptive value size at onset of maturity depends on multitude of exogenous of the allometry exhibited by these organs has already been factors, such as temperature and/or food availability. er Th e- largelydiscussed[6,10,15,34].Inthiscontext,the results fore, it is not a constant character and may vary in different forrelativegrowthinthe P. algeriense female abdomen are as populations of the same species [45]. expected, with the abdomen width still growing during the Marine [15] and freshwater [46, 47] crabs show the well- adult phase. In fact, this phenomenon could provide an extra known, although not well-understood [48], phenomenon space for the xfi ation of eggs on the setose pleopods and thus of cheliped asymmetry. Researchers sometimes seem to act as an incubation chamber for the developing eggs, which discard this asymmetry focusing mainly on investigating the is similar to the general pattern of female abdominal growth functional use of the large cheliped. of most freshwater, marine, and terrestrial crabs [6, 35, 36]. Potamon algeriense is heterochelous, with right- On the other hand, Hartnoll [36] noted that the male handedness in both sexes being the most dominant (89.68% abdomenforms asingleprotectingcover forthegonopods. of males and 89.80% of females). According to Scalici and Finney and Abele [10] are of the opinion that abdomen Gherardi [48], this pattern can be found in most brachyuran width growth in males does not have any reproductive ben- species that present handedness, as previously reported for efits. er Th eby, aer ft the complete gonopod development, the some potamonautid crab species (for details see [47]), as well abdomen growth assumes the isometric, or even a negative, as marine crabs (e.g., [49, 50]). allometrypattern,aspreviouslyobservedforotherfreshwater ec Th helipedof malesislargerthanthatofthe femalesof crabs such as the potamids Potamon potamios palestinensis the same size, suggesting its behavioral utilization, including [33] and the P. u fl viatile [37]; the trichodactylids Dilocarcinus sexual signaling during courtship, intermale fighting for the pagei, Sylviocarcinus australis [38], and Trichodactylus u fl vi- access of females, and the defense of resources such as food atilis [39]. and territories [15, 33, 51–57]. Males with larger cheliped In the literature, besides the abdomen width, there are might increase the likelihood of winning during agonistic frequent citations about the importance of morphometry and interactions for access to receptive females, considering that Log Abdomen Width (mm) Log Abdomen Width (mm) International Journal of Zoology 5 Table 1: Results of allometric relationships studied in Potamon algeriense: analyses of covariance (ANCOVA) carried out to test slope and intercept for both sexes that indicate the beginning of maturity in relation to body size. Significance was indicated by an asterisk. Factor: Relationship Morphometric measure Parameter F Sex/maturity Juveniles vs. Adults Slope 24.157∗ WW Intercept 32.452∗ Juveniles vs. Adults Slope 4.873 CL Intercept 6.486 Juveniles vs. Adults Slope 789.149∗ AW Intercept 848.941∗ Females Juveniles vs. Adults Slope 4.623 AL Intercept 3.453 Juveniles vs. Adults Slope 468.157∗ ChW Intercept 518.614∗ Juveniles vs. Adults Slope 432.181∗ ChL Intercept 495.214∗ Juveniles vs. Adults Slope 3.121 WW Intercept 2.867 Juveniles vs. Adults Slope 12.342 CL Intercept 18.658 Juveniles vs. Adults Slope 2.032 AW Intercept 1.091 Males Juveniles vs. Adults Slope 1.456 AL Intercept 1.017 Juveniles vs. Adults Slope 761.646∗ ChW Intercept 803.558∗ Juveniles vs. Adults Slope 698.154∗ ChL Intercept 645.167∗ Females vs. Males Slope 5.314 WW Intercept 6.185 Females vs. Males Slope 3.194 CL Intercept 4.684 Females vs. Males Slope 64.235∗ AW Intercept 61.489∗ Females/males Females vs. Males Slope 4.135 AL Intercept 5.354 Females vs. Males Slope 82.318∗ ChW Intercept 84.343∗ Females vs. Males Slope 75.156∗ ChL Intercept 76.685∗ ∗P< 0.05 WW: wet weight; CL: carapace length; AW: abdomen width; AL: abdomen length; ChW: cheliped width; ChL: cheliped length. dominance in crabs is largely determined both by body size females with larger cheliped have a better chance of defending (described earlier) and by relative cheliped size [6, 58, 59]. the developing young against attacks from predators [56]. The handedness recorded for females is not a rare situ- ation for freshwater crabs, as it was previously reported for 5. Conclusions some other species such as Candidiopotamon rathbunae in Taiwan [56] and T. fluviatilis [39] and Potamonautes warreni Estimates for the beginning of morphological sexual maturity in South Africa [47]. The use of large cheliped for fights has may provide various results when different body structures also been proposed for ovigerous females. In fact, freshwater are studied, or even when different mathematical methods crabsarewellknowntobedirectdeveloperswithlong are applied. eTh reby, bioassays for mating are suggested incubation periods (for up to three months in Potamonautes for assessing the importance of body parts as a reference sidneyi) and extensive maternal care [51]. Consequently, to estimate the beginning of sexual maturity for crabs, as 6 International Journal of Zoology Table2:Regressionanalysesofmorphometricdataof Potamon algeriense. Carapace width (CW) was used as the independent variable. b 2 ∗ Variable Sex/Stage N Power Equation Y=a.X logY= loga + b logX R t (b=1) Allometry level 2.87 JM 187 WW=9.772CW logWW=2.87 logCW+0.990 0.961 2.91 = 3.17 AM 191 WW=3.890CW logWW=3.17 logCW+0.590 0.988 3.67∗ + WW 2.95 JF 181 WW=5.129CW logWW=2.95 logCW+0.710 0.984 3.05 = 2.81 AF 110 WW=8.710CW logWW=2.81 logCW+0.940 0.957 2.89 = 1.02 JM 187 CL=1.071CW logCL=1.02 logCW+0.029 0.993 1.09 = 1.04 AM 191 CL=1.015CW logCL=1.04 logCW+0.006 0.991 1.14 = CL 0.97 JF 181 CL=0.955CW logCL=0.97 logCW-0.020 0.993 0.94 = 0.90 AF 110 CL=1.230CW logCL=0.90 logCW+0.090 0.982 0.89 = 0.99 JM 187 AW=0.602CW logAW=0.99 logCW-0.220 0.972 1.01 = 0.93 AM 191 AW=0.741CW logAW=0.93 logCW-0.130 0.969 0.72∗ - AW 1.12 JF 181 AW=0.275CW logAW=1.12 logCW-0.560 0.910 1.12 = 1.47 AF 110 AW=0.100CW logAW=1.47 logCW-1.000 0.813 4.65∗ + 1.00 JM 187 AL=0.436CW logAL=1.00 logCW-0.361 0.958 1.00 = 0.89 AM 191 AL=0.602CW logAL=0.89 logCW-0.220 0.927 0.95 = AL 1.19 JF 181 AL=0.309CW logAL=1.19 logCW-0.510 0.939 1.15 = 1.15 AF 110 AL=0.380CW logAL=1.15 logCW-0.420 0.887 1.10 = 1.03 JM 187 ChW=0.741CW logChW=1.03 logCW-0.130 0.926 1.02 = 1.38 AM 191 ChW=0.245CW logChW=1.8 logCW-0.611 0.913 9.89∗ + ChW 1.17 JF 181 ChW=0.182CW logChW=1.17 logCW-0.740 0.908 1.17 = 1.36 AF 110 ChW=0.097CW logChW=1.36 logCW-1.013 0.797 7.34∗ + 1.17 JM 187 ChL=0.213CW logChL=1.17 logCW-0.672 0.952 1.16 = 1.52 AM 191 ChL=0.067CW logChL=1.52 logCW-1.174 0.945 8.56∗ + ChL 1.11 JF 181 ChL=0.501CW logChL=1.11 logCW-0.300 0.962 1.10 = 1.20 AF 110 ChL=0.380CW logChL=1.20 logCW-0.420 0.876 8.20∗ + ∗ t-test;𝛼 =0.05. 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Allometric Growth of the Freshwater Crab Potamon algeriense (Bott, 1967) (Decapoda, Brachyura, Potamidae) in Oued Zegzel, a Mountain Stream, in the Northeast of Morocco

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Hindawi International Journal of Zoology Volume 2019, Article ID 5168639, 8 pages https://doi.org/10.1155/2019/5168639 Research Article Allometric Growth of the Freshwater Crab Potamon algeriense (Bott, 1967) (Decapoda, Brachyura, Potamidae) in Oued Zegzel, a Mountain Stream, in the Northeast of Morocco 1 1 2 1 Soufiane Fadlaoui , Mohammed Mahjoub, Ouahid El Asri, and Mohammed Melhaoui Mohamed First University, Faculty of Sciences, Department of Biology, Laboratory of Water, Environment, and Sustainable Development, P.B. 717, Oujda, Morocco Biochemistry and Biotechnology Laboratory, Mohamed First University, Oujda, Morocco Correspondence should be addressed to Soufiane Fadlaoui; soufiane.fadlaoui@gmail.com Received 30 January 2019; Revised 4 April 2019; Accepted 27 June 2019; Published 1 August 2019 Academic Editor: o Th mas Iliffe Copyright © 2019 Soufiane Fadlaoui et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. A morphometric study has been carried out for the first time on the population of the freshwater crab Potamon algeriense inhabiting Oued Zegzel, a tributary of the Lower Moulouya River, in the northeast of Morocco. Crabs were collected monthly over one year (October 2017 to September 2018) by excavating burrows and searching under blocks. A total of 669 crabs were obtained, 291 females and 378 males. Regression analysis was performed among carapace width (CW), as the reference dimension, and wet weight (WW),carapace length (CL),lengthand widthofabdomen (ALandAW), andlengthand widthofthecheliped (ChL and ChW) were chosen as dependent variables, using the allometric method. Based on Somerton’s technique, the onset of sexual maturity wasestimatedto occurataround27mmCWfor malesand32mmCW forfemales.Thegrowthpatternsrecordedfor P. algeriense are associated with the species reproductive strategy, i.e., preparation of body parts involved in female acquisition and egg incubation, like the male cheliped and the female abdomen, respectively. Among the 291 females sampled, 10.20% were left-handed and 89.80% were right-handed, while among the 378 males examined, 10.32% were left-handed and 89.68% were right-handed. It was concluded that the cheliped width and the abdominal width of P. algeriense are the morphometric variables most appropriate to estimate the size at the beginning of the sexual maturity for males and females of this species, respectively. 1. Introduction and to establish sexual maturity as well as contributing to the identification of “handedness” occurrence [9–12]. Allometry, which is the oldest of the approaches and still Size at which sexual maturity is attained is important widely applied in biology, is a well-known phenomenon in crab life history, because both age and size at maturity concerning the study of the relationship between the size and are associated with a species’ reproductive output [13]. An function of organs of the body and growth or size of the whole assemblage of morphological, physiological, and behavioral body [1]. Allometry is oeft n divided into three principal types: changes through which immature individuals become able ontogenetic (when shape changes with ontogenetic stage or to produce and transfer gametes marks the onset of maturity age), static (when shape correlates with size independently [14]. of age), and evolutionary (when shape correlates with size eTh growth pattern of some specicfi body parts such as among species) [2]. chelipeds, abdomen, and gonopods shows variations in the Relative growth in brachyuran crabs has been commonly degree of allometry during the course of development, which studied since the earlier decades of the century [3–5], but only may coincide with gonad maturation, providing an important in more recent work were these allometric patterns reviewed estimate for the size at which these animals are ready for for the group [6]. It can be used in species identification [7, 8] reproduction [15, 16]. O. Bouizi Oued Cherraa 2 International Journal of Zoology Nador Mediterranean Sea Plain of Triffa W E W E Berkane Midar S S Oujda O. Bou Rdim Aknoul Oued Moulouya Taourirt O. Ouizert O. Charef Taza km O. Charef 0 2 4 Ifrane O. Azreg Azrou Missour Khenifra Midelt Figure 1: Localization of the study area, Oued Zegzel, Morocco. Crabs of the infraorder Brachyura are among the most stream, in the northeast of Morocco, focusing on the mor- diverse groups of crustaceans with more than 7000 described phometric relationships between carapace width and the size species in 98 families, occurring in various habitats: marine, of other body components. Through the analyses of these freshwater, and terrestrial [17–20]. parameters, it is possible to characterize the allometry levels Freshwater crabs are found exclusively in freshwater or and to make inferences about the sizes at which males and terrestrial ecosystems and never enter brackish or marine females reach sexual maturity, thus evidencing their puberty waters for reproduction [21]. Approximatively, 1400 species molt. The frequency of handedness (major cheliped) in each are known with additional species being regularly discovered sex and its growth are also studied. reviewed in [16, 17]. Potamon algeriense belongs to the family of the Potamidae 2. Material and Methods which is the largest of all freshwater crab families and com- 2.1. Study Area. The study was performed in Oued Zegzel prises 95 genera and over 505 species distributed through- ∘ 󸀠 ∘ 󸀠 (34 50 29.4 N, 2 21 19.8 W), a tributary of the Lower out the southern Palaearctic and Oriental zoogeographical Moulouya River, in the northeast of Morocco (Figure 1). eTh regions from Morocco as far east as Japan and as far south region is characterized by a Mediterranean climate. as Indonesia [21, 22]. In fact, it is the only Maghrebian Shrublands that mark the study area were composed representative of this family. eTh species could be found mainly of Nerium oleander, Tamarix africana, Salix pedi- exclusively in three countries: Morocco, Algeria, and Tunisia cillata, Rubus ulmifolius, Rubus ulmifolius,and Crataegus [23]. In Morocco, the species has been reported from the monogyna.Thus,agroup ofsubmerged plants ( Nasturtium north in the watershed of the Oued Laou near Chefchaouen, officinale , Veronica catenata,and Scrophularia aquatica)is from the northeast in watershed of Moulouya, and from individualized on the banks of the Oued, while in the central the Middle Atlas in the Oued Oum er Rbia watershed near waters Potamogeton natans or Potamogeton pectinatus are Khenifra. oen ft overgrown. Despite the wide distribution of P. algeriense,itsoccur- rence drastically declined during the last two decades due to human impacts (sewage, industrial waste, and agricultural 2.2. Crab Sampling and Measurement. The crabs were col- pesticides). Consequently, P. algeriense is included in the lected monthly over one year (October 2017 to September IUCN red list as least concern species. Comprehensive 2018)byexcavatingburrows andsearchingunderblocks.The biological information about this species is incomplete, as crabs could be caught by plunging an arm into the burrow. previous studies have only treated its geographic distribution However, as the depth of the burrow can reach more than 50 and taxonomic status [24]. cm it was not always possible to catch the crabs before they Thisstudy aims toanalyzeforthefirsttimetherelative escaped to the greater depths. Consequently, the individuals growth of P. algeriense, inhabiting Oued Zegzel, a mountain assessed here represent a random sample. O. Melloulou Atlantic Ocean O. Moulouya Oued Zegzel O. Ouizert Oued M. Idriss 󸀠󸀠 󸀠󸀠 International Journal of Zoology 3 ChL Female Male C CW W A A A AW W CL AL C ChW hW AL AW Figure 2: Position of measurements in Potamon algeriense. CW: carapace width; CL: carapace length; ChL: cheliped length; ChW: cheliped width; AW: abdomen width; AL: abdomen length. All measurements were made to the nearest (0.01 mm) 3. Results using a vernier caliper: carapace width (CW) used as the A total of 669 specimens of P. algeriense were collected, and reference dimension, measured at the widest part; carapace their CW sizes ranged from 5.40 to 26.9 mm for juvenile length (CL); abdominal width (AW); abdominal length (AL) males (N=323), from 27 to 60.30 mm CW for adult males (taken from the widest segment); and cheliped width (ChW) (N=55), 11.80 to 31.9 mm for juvenile females (N=158), and and length (ChL). eTh position of these measurements is 32 to 50.10 mm for adult females (N=133). indicated in Figure 2. According to Somerton’s technique [28, 29], the rela- The specimens were also weighted, wet weight (WW), tionships that best indicate the change in the allometric using an analytical pocket balance to the nearest 0.01 g. coefficient between juveniles and adults phase and the dis- Sex determination was based on the morphology of the tinction between male and female growth are CW vs. AW abdomen (considerably wider in females than in males) and for females and CW vs. ChW for males. In fact, the size the number of pleopods. The presence of eggs and juvenile at which 50% of the females reached maturity is 32 mm in the abdomen of females was also recorded. Chelipeds were of CW based on the CW vs. AW relationship. At this size, designated as major or minor by their size. females present a noticeable growth of the abdomen width Only complete and sexed specimens were used in this resultinginanincreaseoftheallometric coecffi ient(1.38). study. On the other hand, the relationship between ChW and CW in males showed a remarkable increase of the cheliped width 2.3. Statistical Analyses. Relative growth was examined fol- soon aer ft the puberty molt, which may occur in crabs of lowing the method of Huxley [25] to describe most patterns approximately 27 mm CW, resulting in an increase of the of relative growth in animals in general and in brachyuran allometric coefficient (1.38) (Figure 3). crabs especially [15, 26, 27]. The regressions parameters (slope and intercept) and the The logarithmic transformation log Y = log a + b log X relative growth equations of the morphometric relationships was linearized from the traditional allometric growth equa- obtained for both sexes in juveniles and adults, and their tion Y = aX , where (Y) and (X) are morphological dimen- departures from isometry (t-test) are shown in Tables 1 and sions and (a) and (b) are growth constraints. eTh computer 2, respectively. programs Mature Iand II[28,29]wereusedtodelimit the er Th elationshipsofWW,AW,ChW,andChLwithCW size at which 50% of the individuals reached maturity. as an independent variable indicate that there are significant Analysis of covariance (ANCOVA) was performed to differences in growth between juveniles and adults in both compare slopes and intercepts of the obtained regressions sexes. within each allometric relationship and between sexes [30, Concerning handedness, right-handed considerably out- 31]. Carapace width (CW) was chosen as the reference numbered left-handed ones; from 669 crabs examined, dimension (independent variable), since this has been widely 10.31% (69 of 669) had the left propodus and dactylus used for recent studies on decapods including the Potamidae enlarged (left-handed) and 89.69% (600 of 669) had the [32]. right propodus and dactylus enlarged (right-handed). No Departures from isometry (Ho: b= 3) for wet weight and homochelous individuals were found. Among the 291 females (Ho: b= 1) for the other measurements were tested using a sampled, 10.20% (30 of 291) were left-handed and 89.80% Student’s t-test on the obtained slope values (𝛼=0.05) [31]. (261 of 291) were right-handed, while among the 378 males If b> 1, then the dimension (Y) increases in size relatively examined, 10.32% (39 of 378) were left-handed and 89.68% more rapidly than the reference dimension (X) does and (339 of 669) were right-handed. No significant difference in growth is said to be positively allometric. A condition of b< handedness between sexes was observed (𝜒 =4.16; P> 0.05). 1 indicates negative allometry, and b = 1 indicates a condition of isometry that means there is no change in the relative 4. Discussion shape with increasing size. Handedness between sexes was analyzed using a two-tailed𝜒 test to determine whether the In crustaceans, some aspects such as gonadal development, frequencies of right- and left-handed males were similar. presence of ovigerous females, and morphometric data are 4 International Journal of Zoology 1,60 Males Females 1,40 Adults 1,40 1,20 1,00 Adults 1,20 Juveniles ,80 Juveniles 1,00 ,60 ,40 ,80 ,20 ,60 ,00 1,00 1,20 1,40 1,60 ,75 1,00 1,25 1,50 1,75 Log Carapace Width (mm) Log Carapace Width (mm) (a) (b) Figure 3: Dispersion points and adjusted curve of the relationships: (a) ChW vs. CW; (b) AW vs. CW for males and females of Potamon algeriense, respectively. used to calculate the mean size at the onset of sexual maturity. growth of cheliped dimensions (generally cheliped width) to In fact, the estimation of puberty size may vary according characterize a possible sexual dimorphism and maturation in to the body part analyzed. In the present study, the mean crabs [6, 34]. size of maturity was calculated according to the relationships The size at the onset of sexual maturity of P. algeriense of ChW vs. CW and AW vs. CW for males and females, from Oued Zegzel was greater in females than in males, respectively. es Th e relationships provide the best tfi of sexual andsimilarresultshavebeenfound inthepopulations of maturity of P. algeriense.Gherardiand Micheli[33]remarked Mithraculus forceps, Mithrax tortugae, and Grapsus adscen- that freshwater crabs mate during the intermolt phase when sionis [40–42]. On the contrary, the onset of sexual maturity the male grabs the female within its chelipeds, pointing out for the populations of T. u fl viatilis and D. pagei [39, 43] from the importance of chelipeds for the reproductive process and south-eastern Brazil occurred at the same size interval, as confirming the choice of this body part as the reference for estimated based on chelipeds and abdomen growth for males the analysis of size at sexual maturity. and females, respectively. Delayed sexual maturity could The patterns of relative growth by abdomen, cheliped, and allow females of P. algeriense to live longer, grow larger, and heterochely in P. algeriense follow those already remarked in therefore have higher rate of fecundity [44]. Furthermore, the most crab species studied so far [6], and the adaptive value size at onset of maturity depends on multitude of exogenous of the allometry exhibited by these organs has already been factors, such as temperature and/or food availability. er Th e- largelydiscussed[6,10,15,34].Inthiscontext,the results fore, it is not a constant character and may vary in different forrelativegrowthinthe P. algeriense female abdomen are as populations of the same species [45]. expected, with the abdomen width still growing during the Marine [15] and freshwater [46, 47] crabs show the well- adult phase. In fact, this phenomenon could provide an extra known, although not well-understood [48], phenomenon space for the xfi ation of eggs on the setose pleopods and thus of cheliped asymmetry. Researchers sometimes seem to act as an incubation chamber for the developing eggs, which discard this asymmetry focusing mainly on investigating the is similar to the general pattern of female abdominal growth functional use of the large cheliped. of most freshwater, marine, and terrestrial crabs [6, 35, 36]. Potamon algeriense is heterochelous, with right- On the other hand, Hartnoll [36] noted that the male handedness in both sexes being the most dominant (89.68% abdomenforms asingleprotectingcover forthegonopods. of males and 89.80% of females). According to Scalici and Finney and Abele [10] are of the opinion that abdomen Gherardi [48], this pattern can be found in most brachyuran width growth in males does not have any reproductive ben- species that present handedness, as previously reported for efits. er Th eby, aer ft the complete gonopod development, the some potamonautid crab species (for details see [47]), as well abdomen growth assumes the isometric, or even a negative, as marine crabs (e.g., [49, 50]). allometrypattern,aspreviouslyobservedforotherfreshwater ec Th helipedof malesislargerthanthatofthe femalesof crabs such as the potamids Potamon potamios palestinensis the same size, suggesting its behavioral utilization, including [33] and the P. u fl viatile [37]; the trichodactylids Dilocarcinus sexual signaling during courtship, intermale fighting for the pagei, Sylviocarcinus australis [38], and Trichodactylus u fl vi- access of females, and the defense of resources such as food atilis [39]. and territories [15, 33, 51–57]. Males with larger cheliped In the literature, besides the abdomen width, there are might increase the likelihood of winning during agonistic frequent citations about the importance of morphometry and interactions for access to receptive females, considering that Log Abdomen Width (mm) Log Abdomen Width (mm) International Journal of Zoology 5 Table 1: Results of allometric relationships studied in Potamon algeriense: analyses of covariance (ANCOVA) carried out to test slope and intercept for both sexes that indicate the beginning of maturity in relation to body size. Significance was indicated by an asterisk. Factor: Relationship Morphometric measure Parameter F Sex/maturity Juveniles vs. Adults Slope 24.157∗ WW Intercept 32.452∗ Juveniles vs. Adults Slope 4.873 CL Intercept 6.486 Juveniles vs. Adults Slope 789.149∗ AW Intercept 848.941∗ Females Juveniles vs. Adults Slope 4.623 AL Intercept 3.453 Juveniles vs. Adults Slope 468.157∗ ChW Intercept 518.614∗ Juveniles vs. Adults Slope 432.181∗ ChL Intercept 495.214∗ Juveniles vs. Adults Slope 3.121 WW Intercept 2.867 Juveniles vs. Adults Slope 12.342 CL Intercept 18.658 Juveniles vs. Adults Slope 2.032 AW Intercept 1.091 Males Juveniles vs. Adults Slope 1.456 AL Intercept 1.017 Juveniles vs. Adults Slope 761.646∗ ChW Intercept 803.558∗ Juveniles vs. Adults Slope 698.154∗ ChL Intercept 645.167∗ Females vs. Males Slope 5.314 WW Intercept 6.185 Females vs. Males Slope 3.194 CL Intercept 4.684 Females vs. Males Slope 64.235∗ AW Intercept 61.489∗ Females/males Females vs. Males Slope 4.135 AL Intercept 5.354 Females vs. Males Slope 82.318∗ ChW Intercept 84.343∗ Females vs. Males Slope 75.156∗ ChL Intercept 76.685∗ ∗P< 0.05 WW: wet weight; CL: carapace length; AW: abdomen width; AL: abdomen length; ChW: cheliped width; ChL: cheliped length. dominance in crabs is largely determined both by body size females with larger cheliped have a better chance of defending (described earlier) and by relative cheliped size [6, 58, 59]. the developing young against attacks from predators [56]. The handedness recorded for females is not a rare situ- ation for freshwater crabs, as it was previously reported for 5. Conclusions some other species such as Candidiopotamon rathbunae in Taiwan [56] and T. fluviatilis [39] and Potamonautes warreni Estimates for the beginning of morphological sexual maturity in South Africa [47]. The use of large cheliped for fights has may provide various results when different body structures also been proposed for ovigerous females. In fact, freshwater are studied, or even when different mathematical methods crabsarewellknowntobedirectdeveloperswithlong are applied. eTh reby, bioassays for mating are suggested incubation periods (for up to three months in Potamonautes for assessing the importance of body parts as a reference sidneyi) and extensive maternal care [51]. Consequently, to estimate the beginning of sexual maturity for crabs, as 6 International Journal of Zoology Table2:Regressionanalysesofmorphometricdataof Potamon algeriense. Carapace width (CW) was used as the independent variable. b 2 ∗ Variable Sex/Stage N Power Equation Y=a.X logY= loga + b logX R t (b=1) Allometry level 2.87 JM 187 WW=9.772CW logWW=2.87 logCW+0.990 0.961 2.91 = 3.17 AM 191 WW=3.890CW logWW=3.17 logCW+0.590 0.988 3.67∗ + WW 2.95 JF 181 WW=5.129CW logWW=2.95 logCW+0.710 0.984 3.05 = 2.81 AF 110 WW=8.710CW logWW=2.81 logCW+0.940 0.957 2.89 = 1.02 JM 187 CL=1.071CW logCL=1.02 logCW+0.029 0.993 1.09 = 1.04 AM 191 CL=1.015CW logCL=1.04 logCW+0.006 0.991 1.14 = CL 0.97 JF 181 CL=0.955CW logCL=0.97 logCW-0.020 0.993 0.94 = 0.90 AF 110 CL=1.230CW logCL=0.90 logCW+0.090 0.982 0.89 = 0.99 JM 187 AW=0.602CW logAW=0.99 logCW-0.220 0.972 1.01 = 0.93 AM 191 AW=0.741CW logAW=0.93 logCW-0.130 0.969 0.72∗ - AW 1.12 JF 181 AW=0.275CW logAW=1.12 logCW-0.560 0.910 1.12 = 1.47 AF 110 AW=0.100CW logAW=1.47 logCW-1.000 0.813 4.65∗ + 1.00 JM 187 AL=0.436CW logAL=1.00 logCW-0.361 0.958 1.00 = 0.89 AM 191 AL=0.602CW logAL=0.89 logCW-0.220 0.927 0.95 = AL 1.19 JF 181 AL=0.309CW logAL=1.19 logCW-0.510 0.939 1.15 = 1.15 AF 110 AL=0.380CW logAL=1.15 logCW-0.420 0.887 1.10 = 1.03 JM 187 ChW=0.741CW logChW=1.03 logCW-0.130 0.926 1.02 = 1.38 AM 191 ChW=0.245CW logChW=1.8 logCW-0.611 0.913 9.89∗ + ChW 1.17 JF 181 ChW=0.182CW logChW=1.17 logCW-0.740 0.908 1.17 = 1.36 AF 110 ChW=0.097CW logChW=1.36 logCW-1.013 0.797 7.34∗ + 1.17 JM 187 ChL=0.213CW logChL=1.17 logCW-0.672 0.952 1.16 = 1.52 AM 191 ChL=0.067CW logChL=1.52 logCW-1.174 0.945 8.56∗ + ChL 1.11 JF 181 ChL=0.501CW logChL=1.11 logCW-0.300 0.962 1.10 = 1.20 AF 110 ChL=0.380CW logChL=1.20 logCW-0.420 0.876 8.20∗ + ∗ t-test;𝛼 =0.05. 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