crustacean research, no. 40: 13 – 20, 2011 13 A rare macrophthalmine , Euplax leptophthalmus H. Milne Edwards, 1852 (, Brachyura, ) from Amami-Oshima Island, Ryukyu Islands, southern Japan

tei Kishino, toshihiko Yonezawa and Keiji Wada

Abstract.– The rare macrophthalmine of E. leptophthalmus and M. gastrodes, and crab, Euplax leptophthalmus H. Milne concluded that both species were identical, Edwards, 1852, is recorded from Japan except in a few details of the male chelae. for the first time. Numerous specimens as for the , Barnes (1966) had were obtained from tidal muddy mangrove synonymised Euplax under creeks on Amami-Oshima Island. Variation Desmarest, 1823. Barnes (1967) recognized is discussed with regard to sexual and size six subgenera under Macrophthalmus in differences. Differences with the allied E. his revision, but inexplicably placed M. dagohoyi (Mendoza & Ng, 2007) from the leptophthalmus in his new subgenus Venitus Philippines are also discussed. Barnes, 1967 (type species Gonoplax latreillei Desmarest, 1822), and chose not to recognize Euplax h. Milne edwards, 1852, even though Introduction it was an available name and had precedence t he brachyuran subfamily (see Mendoza & ng, 2007). Barnes (1977) Macrophthalminae Dana, 1851 (family commented that M. gastrodes was a junior Macrophthalmidae Dana, 1851) currently synonym of M. (Venitus) leptophthalmus has 9 genera, 3 subgenera and 61 species with the former considered a juvenile form (ng et al., 2008; McLay et al., 2010). on of the latter and noted that “chile”, the type the Japanese coast, 23 species are known in locality of M. leptophthalmus, was erroneous. three genera: of Macrophthalmus Desmarest, Mendoza & ng (2007) agreed that the type 1823, Chaenostoma stimpson, 1858 and locality of Euplax leptophthalmus was clearly Venitus Barnes, 1967 (sakai, 1976; Kosuge & erroneous and suggested that a site in the Davie, 2001; sakai, 2003; nagai et al., 2006; Indian ocean may be the type locality instead. naruse & Kosuge, 2008). however, the genus recently, Mendoza & ng (2007) revised Euplax h. Milne edwards, 1852 has not been the identity of M. leptophthalmus and recorded from Japan. resurrected the Euplax as a separate subgenus Euplax was established by h. Milne of Macrophthalmus. they also showed that edwards (1852) for a new species Euplax the subgenus Venitus could be separated from leptophthalmus, described from one male and Euplax. In addition, a new species of Euplax, one female specimen collected in “chile”, M. (E.) dagohoyi, was described from the south america. Kemp (1915) later described Philippines. recently, McLay et al. (2010) a closely allied species, Macrophthalmus regarded both Euplax and Venitus as full gastrodes from the chilka Lake in India. genera. Kemp (1915), however, did not compare M. We collected a good series of male and gastrodes and E. leptophthalmus. Barnes female macrophthalmine that had not (1977) re-examined morphological characters been reported in Japan from tidal muddy 1414 t. KIshIno Et al.

mangrove creeks in amami-oshima Island of southern Japan from 2000 to 2004. these specimens were eventually identified with Euplax leptophthalmus h. Milne edwards, 1852. For E. leptophthalmus, only four specimens have been found thus far and not much is known about variation in the taxon. In this paper, we show that amami-oshima Island specimens are true E. leptophthalmus. the good series of specimens obtained also allow us to comment on morphological variations in the species associated with size and sex.

Materials and Methods specimens were collected by a hand net having mesh size 2 mm, from mangrove creeks (water depth: 20–100 cm at low tide) at the Yakugachi river estuary (28˚15’4”n, 129˚24’30”e), tekebu Mangrove swamp (28˚26’24”n, 129˚40’0”e) and the urakawa river estuary (28˚24’57”N, 129˚36’2”E) in amami-oshima Island, mid-ryukyu Islands of southern Japan in 2000 to 2004. these 15 specimens were preserved in 80% ethanol and deposited in the osaka Museum of Fig. 1. Euplax leptophthalmus. Male (14.0 by 11.4 national history in osaka, Japan (oMnh- mm) (oMnh-ar8697). a, dorsal view; B, ar). the specimens, entirely covered with ventral view; c, frontal view. mud, were washed and observed under a microscope of 8–40 magnifications. Staining was made with methylene blue as and when pl. XII, fig. 5. necessary. carapace size is expressed as cW Macrophthalmus leptophthalmus: Barnes, 1966: pl. (carapace width) by cL (carapace length), XXIV, figs. 3, 4. with cW measured at the widest level. other Macrophthalmus (Venitus) leptophthalmus: Barnes, measurements followed Komai et al. (1995: 1977: text-fig. 1. fig. 1). Macrophthalmus (Euplax) leptophthalmus: Mendoza & Ng, 2007: text-figs 1, 2; Ng et al., 2008: 237 (list); Davie, 2009: 820 (key); Barnes, 2010: text- systematics fig. 1b (as schematic illustration). Macrophthalmidae Dana, 1851 Material examined.– 1 male (14.0 by Macrophthalminae Dana, 1851 11.4 mm) (oMnh-ar8697), subtidal mud in Euplax h. Milne edwards, 1852 mangrove creek, Yakugachi river estuary, Euplax leptophthalmus h. Milne edwards, amami-oshima Island, 11 May 2002, 1852 collected by t. Yonezawa; 1 female (18.0 by (Figs. 1–3) 14.8 mm) (oMnh-ar8696), subtidal mud in mangrove creek, Yakugachi river estuary, 6 Euplax leptophthalmus h. Milne edwards, 1852: 160 (type locality: chile?, south america); rathbun, nov. 2001, coll. t. Yonezawa; 1 male (11.1 1918: 423. by 8.9 mm = exuvium size just after molting) Macrophthalmus gastrodes Kemp, 1915: (type (oMnh-ar8695), subtidal mud in mangrove locality: the Chilka Lake, India), text-figs. 9, 10. creek, Yakugachi river estuary, 4 June 2000, Euplax lEptophthalMus FroM JaPan 1515

Fig. 2. Euplax leptophthalmus. Male (14.0 by 11.4 mm) (oMnh-ar8697): a, c–G; female (18.0 by 14.8 mm) (oMnh-ar8696): B. a, carapace (dorsal view); B, female antero-lateral tooth of left side (dorsal view); c, frontal view of left side; D, right third maxilliped (ventral view); e, right 2nd – 4th ambulatory legs (dorsal view); F, left first gonopod (ventral view); G, distal part of left first gonopod (ventral view). Scales, A– F = 5 mm; G = 3 mm. coll. t. Kishino; 2 males (9.0 by 8.0 mm, equal to carapace length; regions well defined, 10.3 by 9.1 mm), 3 females (7.8 by 6.8 mm to with 2 deep longitudinal grooves marking 11.3 by 9.7 mm), 1 juvenile (4.2 by 3.7 mm) both sides of the gastric and cardiac regions; (oMnh-ar8698), subtidal mud in creek, surface with scattered very short setae, with tekebu, amami-oshima Island, 6 nov. 2003, patches of granules on gastric, cardiac and coll. t. Yonezawa and t. Kishino; 2 males branchial regions. Lateral margins convergent (6.7 by 5.5 mm, 6.9 by 5.6 mm) (oMnh- anteriorly and posteriorly, point of greatest ar8699), subtidal mud in creek, tekebu, carapace width at postero-lateral angle (at amami-oshima Island, 16 nov. 2003, coll. t. level of second/third ambulatory legs). Kishino; 1 male (13.0 by 11.3 mm) (oMnh- antero-lateral margin granulated, setose, with ar8700), subtidal mud in mangrove creek, three well-defined (including orbital corner) urakawa river estuary, 5 apr. 2004, coll. small teeth and inconspicuous fourth tooth. t. Yonezawa; 2 males (9.8 by 9.1 mm, 12.2 First (orbital corner) and second tooth broad, by 10.5 mm), 1 female (11.6 by 10.1 mm) subtriangular, bluntly pointed; third tooth (oMnh-ar8701), subtidal mud in creek, smaller than former tooth, bluntly triangular, tekebu, amami-oshima Island, 6 apr. 2004, fourth present as obscurely granulated coll. t. Yonezawa. protrusion. Postero-lateral margin and Description of male (oMnh-ar8697).– posterior margin smooth excepting inferior carapace subquadrate (Figs. 1a; 2a), almost ridge lined by very small granules. circular, in shape, width 1.2 times of length; Front (Figs. 1a, c; 2a, c) moderate in width across external orbital angles almost width (1/5 carapace width at external orbital 1616 t. KIshIno Et al.

Fig. 3. Euplax leptophthalmus. Male (14.0 by 11.4 mm) (oMnh-ar8697): left sides; female (18.0 by 14.8 mm) (oMnh-ar8696): right sides. a, B: abdomen and left side of thoracic sternites; c, e: left chela (outer view); D, F: left chela (inner view). scales, 5 mm.

angle), slightly constricted between bases of septum; basal segment distinctly inflated. eyestalk; frontal margin granulated, deep, antennae almost as long as eyestalks. surface without granules, anterior margin epistome (Figs. 1c; 2c) narrow, central slightly emarginated medially by longitudinal region of posterior border of epistome groove. eyestalks prominent, corneas concave, without posterior rim or postero- reduced, cornea only extending to a point median protrusion; endostominal ridge just short of base of external orbital angle. in anterior buccal cavity absent. third upper orbital margin curved, slightly sloping maxillipeds separated by a median hiatus, backwards; margin granulated, setose. Lower setose along internal margins; merus and orbital margin granulated, lined by long setae. ischium with longitudinal sculpturing near antennules separated by a narrow median inner margin; merus about half as long as Euplax lEptophthalMus FroM JaPan 1717 ischium; internal and external margins of abdomen (Figs. 1B; 3a) with 6 freely merus convex; anterior margin excavated; articulating somites and telson with short internal margin of ischium slightly concave setae along margins, surface almost smooth, medially, external margin almost straight, central convexities developed; 1st somite posterior margin with small rounded granules; short, widest with obscure transverse ridge exopod narrow, width about 0.3 times of bearing minute granules in median surface; ischium, with flagellum. 2nd somite shortest, narrower than 1st and chelipeds subequal (Fig. 1a); cheliped 3rd somites; 3rd somoite trapezoidal, slightly length shorter than 1 st, longer than 4 th narrower than 1st somite; 4th somite to telson ambulatory legs. surface of merus and carpus tapering gradually; telson width (at base) almost smooth without spine or crenulate subequal to length, tip rounded. surface of tooth. Merus with row of long setae on upper sternites almost smooth excepting border with and lower margins; carpal margin with long telson and 6th somite bearing row of granules; setae; inner and outer surfaces with scattered space from anterior margin of telson to short setae. Inner surface of carpus covered posterior margin of buccal cavity extremely with long setae, other surfaces same as short. merus; joint with palm lined sparsely by long First pleopod (Fig. 2F, G) almost straight, setae. Palm (Fig. 3c, D) length subequal to subdistal part tapering; terminal process fingers, slightly inflated; single row of long curved outwards, tip sharp; external margin setae lining upper margin, median external and distal end of inner margin with a row surface and lower margin, continuing to of setae from ventral view; distal part and fixed finger from outer view; inner surface terminal process concealed beneath dense concealed beneath thick mat of plumose long setae. setae excepting smooth surface near lower Description of female (oMnh-ar8696).– margin and near joint with carpus. Fixed carapace, orbits, epistome, exognath and finger almost straight; row of setae near ambulatory legs very similar to male. cutting margin on inner and outer sides; with surface of carapace relatively more large crenulated, quadrangular tooth medially. granular than male, granulated area occupying Dactylus slightly curved; row of setae lining about half of carapace; 1st tooth on antero- upper margin; row of setae lining both lateral margin blunter than male (Fig. 2B); outer and inner cutting margins from large entire margin bearing minute granules. crenulated tooth to tip; with a differentiated, chelipeds subequal, small, slender; length molariform tooth on proximal cutting margin. shorter than 4th ambulatory leg; setae of ambulatory legs (Figs. 1a; 2e) long, margins and surfaces almost similar to male. slender, not granulose on surface; 3rd legs Inner surface of palm (Fig. 3e, F) thinly longest, about twice as long as carapace setose with a row of setae on upper lateral width. Merus length subequal to combined region. Both fingers nearly straight, sharp; carpus and propodus lengths; anterior margin cutting margin without distinct tooth as in with inconspicuous crenulate tooth and small male, fixed finger with weakly crenulated subterminal spine except on 4th legs; posterior tooth, dactylus almost smooth. ambulatory margin with obscure crenulate tooth except legs similar to male except in presence of 3rd and 4th legs; long setae lining both anterior crenulated tooth on posterior margin of merus and posterior margins. carpi and propodi of of 3rd leg. 1st to 3rd legs with thick mats of short plumose abdomen (Fig. 3B) wide, covering most setae across anterior lateral surfaces to upper of thoracic sternites; 6 freely articulating margins; lower margins with long setae. somites and telson with marginal long setae, Dactyli of 1st to 3rd legs, flattened, narrowly surface almost smooth, central convexities straight lanceolate, anterior margins covered well developed; 1st somite shortest, obscure with short setae; 4th dactyli slightly curved transverse ridge structured by minute granules outwards, all margin with long setae; a row of in antero-lateral surface; 4th and 5th somites setae on longitudinally median portion in all widest; telson width (at base) slightly longer dactyli. than length, tip rounded anteriorly. surface of 1818 t. KIshIno Et al.

sternites (Fig. 3B) almost smooth excepting mythology, because the reduced corneas, border with telson bearing a row of granules; whitish body coloration and whole body space from anterior margin of telson to wearing with mud, look like a ghost in hades. posterior margin of buccal cavity extremely Remarks.– the specimens collected in short. amami-oshima Island can be undoubtedly Morphological variations by size and placed in Euplax h. Milne edwards, 1852 sex.– Mean (ranges) of cW/cL ratio in male by the following features: 1) carapace less = 1.2 (1.1–1.2, N = 9), in female = 1.2 (1.1– than 1.2 times as wide as long, front slightly 1.2, N = 5), in juvenile = 1.1 (N = 1). constricted at base of eyestalks, external the area of granules on carapace orbital angle small and blunt (Figs. 1a; surface changed with body size, with larger 2a, B); 2) lower orbital border with small individuals having proportionately granules. granules, epistome narrow, postero-median this was supported by statistics (spearman’s margin of epistome broadly concave, without correlation coefficient by rank test: N = 14 rim on posterior margin (Figs. 1c; 2c); (excepting a molting individual), p<0.01, z = 3) third maxilliped with merus distinctly 2.65). shorter than ischium (Figs. 1B, c; 2D); and the possession of blunter 1st antero-lateral 4) dactylus of male chela with differentiated tooth (Fig. 1B) was not diagnostic for females tooth on the cutting margin (Fig. 3c, D). because two male specimens (9.8 mm cW, the carapace, orbits, chelae, ambulatory 12.2 mm cW) had the same condition. the legs and abdomen of amami-oshima 4th antero-lateral tooth was present in all specimens agreed well with those of E. specimens, with a juvenile (4.2 mm cW) leptophthalmus as defined by Barnes (1977) having a distinct one. and Mendoza & ng (2007). Furthermore, the male cheliped changed with body according to Barnes (1977), the male first size, with small specimens having relatively gonopod of E. leptophthalmus is more or less more slender and smaller chelipeds with straight, with moderate developed, curved sparse setae on the inner surface of palm as terminal process. this description of first in females (Fig. 3E, F). The male finger also gonopod matches that of the amami-oshima varied by size. only two specimens (12.2 mm specimens. cW, 14.0 mm cW) had a tooth on cutting the carapace, chelipeds, abdomen and margin as in Fig. 3c, D. In contrast, small thoracic sternites of the amami-oshima male specimens lacked the tooth on the finger specimens are relatively less granular than likes female. however, one large male (13.0 those of known E. leptophthalmus (Kemp, mm cW) also lacked the tooth. 1915; Barnes, 1977; Mendoza & ng, 2007). Minute serrate teeth on posterior margin however, the degree of granularity varies of merus of 3rd ambulatory leg were present in with body size, such that larger crabs are some specimens (7/14), while absent in other relatively more granular, a pattern observed in male and female specimens large and small. the amami-oshima specimens. the body size The first gonopod was different in smaller of the known E. leptophthalmus ranged from males: the terminal process was relatively 17.6 to 24.0 mm cW (Kemp, 1915; Barnes, shorter and blunter in smaller specimens 1977), while the largest size of amami- (the smallest male, 6.7 mm cW, lacked a oshima specimen is 14.0 mm cW in male terminal process), and the setae on the inner and 18.0 mm cW in female. It is, therefore, margin from ventral view were also much conceivable that the amami-oshima sparser. Females 7.8 mm and 9.0 mm cW specimens are less granular than the known E. with undeveloped vulvae had abdomens leptophthalmus because they are smaller. which were relatively narrower than those in the genus Euplax currently contains a mature 11.3 mm cW specimen. two species of E. leptophthalmus h. Milne live coloration.– the whole body, after edwards, 1852 and E. dagohoyi (Mendoza removing the mud, was purple whitish. & ng, 2007) (Mendoza & ng, 2007; ng Japanese name.– Yomi-no-osagani. et al., 2008; Barnes, 2010). according to “Yomi” in Japanese means hades in Greek Mendoza & ng (2007), Euplax dagohoyi Euplax lEptophthalMus FroM JaPan 1919 differs significantly from E. leptophthalmus E. dagohoyi (see Mendoza & Ng., 2007: fig. in the following features: 1) the carapace 5D, F, G); but almost straight, subdistal part is relatively less granular; 2) the upper tapering, terminal process sharp pointed in E. orbital border leading to and including the leptophthalmus (Fig. 2F, G). first antero-lateral tooth is more backward- some morphological characteristics sloping; 3) the u-shaped notch between (especially, the reduced cornea and lack of the first and second antero-lateral teeth is pigmentation on the body) in Euplax are not as pronounced; 4) the male chelae are related to their habit of hiding in muddy proportionately much smaller, the palm is not riverbed of estuaries. With such a habit, it inflated; instead of a truncated tooth, there is unlikely that Euplax will be frequently is only a ridge on the fixed finger, and the collected, which may be responsible for their gape and inner surface of the palm are only limited records (chilka Lake in India, Bohol sparsely setose (versus densely setose); 5) Island and Panglao Island in Philippines and the ambulatory legs are relatively longer and amami-oshima Island in Japan). We believe more slender; and 6) the sternal rim bordering Euplax is actually more common in muddy the telson and the penultimate abdominal mangrove estuaries from southeast asia to segment is smooth (versus granular). the ryukyu Islands. however, the shape of the notch (u- or V-shape) between the 1st and 2nd antero- acknowledgements lateral tooth may be variable- the shape of the notch of small E. leptophthalmus described We express our thanks to Prof. colin by Kemp (1915) is similar to those of E. McLay, Prof. r. s. K. Barnes and Prof. P. K. L. dagohoyi. the notch shapes of the amami- ng for their reviewing of the manuscript. oshima specimens were also similar to the specimen in Kemp (1915), and the distinct Literature cited u-shape notch as observed in Milne edwards (1852) was not found in either of our all Barnes, r. s. K., 1966. the status of the crab specimens (see Kemp, 1915: fig. 9a, b; genus Euplax h. Milne edwards, 1852; Barnes, 1977: fig. 1b; Mendoza & Ng, 2007: and a new genus australoplax of the figs. 1A, 2A, 3A, 4A; present study: Fig. 2A, subfamily Macrophthalminae Dana, 1851 B). consequently, both species of same body (Brachyura: ). the australian size are likely to have the similar shape of Zoologist, 13: 370–376, including plate the notch. the degree of granulation on the 24. carapace and shape of male chelae are also ———, 1967. the Macrophthalminae of probably variable with body size, especially australasia; with a review of the evolution for juvenile. hence, juvenile forms of the two and morphological diversity of the type species are difficult to distinguish. We believe genus Macrophthalmus (crustacea: that the most useful character indicated by Brachyura). transactions of the Zoological Mendoza & ng (2007) to distinguish the society of London, 31: 195–262. two species that is size-independent is the ———, 1977. concluding contribution sternal rim bordering the telson tip, because towards a revision of, and a key to, the it does not change with sex and body size genus Macrophthalmus (crustacea: in the amami-oshima specimens: smooth Brachyura). Journal of Zoology, London, and long in E. dagohoyi (see Mendoza & 182: 267–280. ng, 2007: figs. 3B, 5a) but granular and ———, 2010. a review of the sentinel short in E. leptophthalmus (see Barnes, and allied crabs (crustacea: Brachyura: 1966: plate 24-fig. 4; Mendoza & Ng, 2007: Macrophthalmidae), with particular fig. 1B; present study: Figs. 1c, 3a, B). In reference to the genus Macrophthalmus. addition, the two species are distinguished Raffles Bulletin of Zoology, 58: 31–49. by the structure of the male first pleopod: Dana, J. D., 1851. conspectus crustacearum slightly curved outwards, subdistal part not quae in orbis terrarum circumnavigatione, tapering, tip of terminal process truncated in carolo Wilkes e classe reipublicae 2020 t. KIshIno Et al.

Foederatae Duce, lexit et descriptsit J. classification naturelle des crustace´s. D. Dana. Proceedings of the academy of annales des sciences naturelles, 18: 109– natural sciences of Philadelphia, 5: 247– 166. 254. nagai, t., Watanabe, t., & naruse, t., 2006. Davie, P. J. F., 2009. a new genus and Macrophthalmus (Macrophthalmus) species of Macrophthalmidae (Brachyura, microfylacas, a new species of sentinel ) from the northern territory, crab (Decapoda: Brachyura: ocypodidae) australia. crustaceana, 82: 815–827. from western Japan. Zootaxa, (1171): Desmarest, a. G., 1822. Les crustacés dits. 1–16. In: a. Brongniart & a. G. Desmarest, naruse, t., & Kosuge, t., 2008. a new (ed), histoire naturelle des crustacés species of Macrophthalmus (crustacea: fossils sous les repports zoologiques et Decapoda: Brachyura: Macrophthalmidae) géologiques, vii + 446pp., Levraut, Paris. from Iriomote Island, ryukyu Islands, ———, 1823. Dictionnaire des sciences Japan. species Diversity, 13: 117–122. naturelles, Vol. 28. F. G. Levreault, Paris. ng, P. K. L., Guinot, D., & Davie, P. J. F., pp. 138–425. 2008. systema Brachyurorum: Part 1. an Kemp, s., 1915. Fauna of the chilka Lake no annotated checklist of extant brachyuran 3. crustacea Decapoda. Memoirs of the crabs of the world. raffles Bulletin of Indian Museum, 5: 199–325. Zoology, supplement (17): 1–286. (With Komai, t., Goshima, s., & Murai, M., 1995. corrigenda and errata parts 1 and 5). crabs of the genus Macrophthalmus of rathbun, M. J., 1918. the grapsoid crabs of Phuket, thailand (crustacea: Brachyura: america. Bulletin of the united states ocypodidae). Bulletin of Marine science, national Museum, 97: 1–461. 56: 103–149. sakai, K., 2003. the Japanese names for the Kosuge, t., & Davie, P. J. F., 2001. Japanese crabs. taxa, (15): 13–30. (In redescription of Macrophthalmus Japanese with english abstract). boteltobagoe and M. holthuisi with notes sakai, t., 1976. crabs of Japan and the on their ecology (Brachyura: ocypodidae). adjacent seas. Kodansha, tokyo. (In 3 Journal of Biology, 21: 545– volumes: (1) english text, xxx + 773 pp., 555. (2) Plate volume, 16 pp., 256 pls, (3) McLay, c. L., Kitaura, J., & Wada, K., 2010. Japanese text, 461 pp.). Behavioural and molecular evidence for the systematic position of Macrophthalmus ( h e m i p l a x ) h i r t i p e s h o m b r o n & Jacquinot, 1846, with comments on addresses: (tK) office of river macrophthalmine subgenera (Decapoda, ecological research: Kaseyama-touda 72-5, Brachyura, Macrophthalmidae). Kizugawa 619-0211, Japan; (tY) Foundation crustaceana Monographs, 14: 483–503. of Kagoshima environmental research and Mendoza, J. c. e., & ng, P. K. L., 2007. service, nanatsujima 1-1-5, Kagoshima 891- Macrophthalmus (Euplax) h. Milne 0132, Japan; (KW) nara Women’s university: edwards, 1852, a valid subgenus of Kita-uoya-nishimachi, nara 630-8506, Japan. ocypodoid crab (Decapoda: Brachyura: e-mails: (tK) [email protected]; Macrophthalmidae), with description of a (tY) [email protected]; (KW) new species from the Philippines. Journal [email protected] of crustacean Biology, 27: 670–680. Milne edwards, h., 1852. observations Received: 8 april 2011. sur les affinities zoologiques et la Accepted: 5 July 2011.