Pagurus rubrior, Komai, 2003

Komai, Tomoyuki, 2003, Identities of Pagurus japonicus (Stimpson, 1858), P. similis (Ortmann, 1892) and P. barbatus (Ortmann, 1892), with description of a new species (Crustacea, Decapoda, Anomura, Paguridae), Zoosystema 25 (3), pp. 377-411 : 401-409

Pagurus rubrior n. sp.

( Figs 6B ; 12 ; 13 )

Eupagurus japonicus – Ortmann 1892: 309, pl. 12, fig. 16. Non Eupagurus japonicus Stimpson, 1858 . See Remarks.

Pagurus similis – Miyake 1960: 90 (part), pl. 45, fig. 5; 1975: pl. 115, figs 6, 9; 1978: 103 (part), fig. 40, pl. 2, fig. 3; 1982: 125 (part), pl. 42, fig. 2; 1991: 125 (part), pl. 42, fig. 2; 1998: 125 (part), pl. 42, fig. 2. — Suzuki 1971: 97, pl. 34, fig. 4. — Kim 1973: pl. 7, fig. 39. — Miyake & Imafuku 1980: 60 (part). — Takeda 1982: 67 (part); 1986: 124, unnumbered fig.; 1994: 228, fig. 5. — Asakura 1995: 362, pl. 97, fig. 4. — Kobayashi 2000: 186, unnumbered fig. — Minemizu 2000: 149, unnumbered fig. — Park & Choi 2001: 139, unnumbered fig. See Remarks.

TYPE MATERIAL. — Holotype: Japan. Hota , Boso Peninsula, lobster net, 10 m, 18.VI.1998, coll. T. Komai, sl 13.2 mm ( CBM-ZC 4778 ).

Paratypes: Japan. Tokyo Bay , exact location unknown, 1882, coll. L. Döderlein, 3 sl 18.9-19.0 mm, 3 sl 14.7-15.7 mm ( MZS 481 ) ; Hota, Boso Peninsula , lobster net, 10-20 m, 21. V.1994, coll. T. Komai, 2 sl 10.7, 16.0 mm, 1 sl 15.1 mm ( CBM-ZC 470 ) ; off Takeoka, Boso Peninsula , gill net, 20-30 m, 1.VIII.1994, coll. T. Komai, 1 sl 13.9 mm, 1 sl 15.9 mm ( CBM-ZC 566 ) ; Takeoka , lobster net, 10-20 m, 29.VIII.1994, coll. T. Komai, 3 sl 10.3- 19.3 mm, 1 sl 12.4 mm, 3 ovig. sl 10.4-16.7 mm ( CBM-ZC 608 ) ; Ubara, Katsuura, Boso Peninsula , scuba diving, 3-6 m, VI.1994, coll. M. Aizawa, 1 sl 11.7 mm ( CBM-ZC 892 ) ; off Takeoka , gill net, c. 30 m, 28.VIII.1996, 1 sl 13.1 mm ( CBM-ZC 2968 ) ; Hota , lobster net, c. 10 m, 18. VI.1998, coll. T. Komai, 3

sl 16.0-21.0 mm ( CBM-ZC 4779); off Takeoka, gill net, 20-30 m, 18. VI.1998, 1 sl 17.3 mm (CBM- ZC 4780); Hota, lobster net, 5-6 m, 20.XII.1998, 2 sl 12.3 mm, 1 sl 17.6 mm ( CBM-ZC 4848); Takeoka, lobster net, 5-10 m, II.1999, 3 sl 11.5- 18.7 mm, 1 (sl 11.7 mm) ( CBM-ZC 6288); similar locality, lobster net, 23.VIII.2000, 1 sl 16.0 mm, 1 ovig. sl 17.6 mm ( CBM-ZC 6289), 1

sl 14.9 mm (MNHN-Pg 6099); similar locality, lobster net, VIII.1997, 1 sl 21.2 mm, 1 ovig. sl 14.3 mm (MNHN-Pg 6098); similar locality, lobster net, 5 m, 8.III.2002, 4 sl 12.1-14.7 mm (MNHN-Pg). — Sagami Bay , exact location unknown, 1903, coll. A. Haberer, 1 sl 14.7 mm, identified by Balss (1913) as Eupagurus japonicus ( ZSM 289 /1) ; Kamegisho , dredge, 14 m, 25.VII.1957, identified by Miyake (1978) as P. similis , det. No. 191, 1 not measured (NSMT- CrR 1372) ; Kamegisho , dredge, 13-14 m, 11.VII.1962, identified by Miyake (1978) as P. similis , det. No. 463, 1 sl 13.5 mm (NSMT-CrR 2012) ; off Hayama, Miura Peninsula , lobster net, 5-10 m, V.1993, coll. H. Ikeda, 1 sl 16.1 mm, 1 sl 16.4 mm (HSM-Cra 0132) ; Kaneda Bay, Miura Peninsula , 10 m, V.1993, coll. H. Ikeda, 1 18.0 mm ( HSM) ; Jogashima Islet, lobster net, 8 m, VIII.1993, coll. H. Ikeda, 2 sl 13.3, 17.0 mm ( HSM). — Kii Peninsula, Izumo , Kushimoto , lobster net, depth not recorded, 23.XI.1976, coll. M. Imafuku, reported by Miyake & Imafuku (1980) as P. similis , 2 sl 6.7, 9.9 mm (OMNH-Ar 1798, 1799) ; off Minabe, lobster net, depth not recorded, 17.XI.1976, coll. M. Imafuku, reported by Miyake & Imafuku (1980) as P. similis , 1 sl 10.6 mm (OMNH-Ar 1826). — Sea of Japan, Takasa, Echizen , Fukui Prefecture, scuba, 10 m, 29. V.2001, coll. T. Sugimoto, 1 sl 15.4 mm, 1 sl 9.4 mm ( CBM-ZC 6447 ) .

TYPE LOCALITY. — Hota, Boso Peninsula, central Japan, at depth of 10 m.

ETYMOLOGY. — The name is derived from the comparative of the Latin adjective ruber and reflects the redder color of this new species compared to the other close relatives of Pagurus .

DISTRIBUTION. — Pacific coast of Japan southward from Boso Peninsula to Kyushu, Sea of Japan coast of southern part of Honshu mainland, and Korea.

HABITAT. — Rocky bottom, subtidal depth to about 30 m. Using gastropod shells, e.g., Turbo cornutus Lightfoot, 1786 , Tonna luteostoma (Küster, 1857) , Cymatium parthenopeum (Salis Marshlins, 1793) , and Charonia lampas sauliae (Reeve, 1844) .

DESCRIPTION

Shield ( Fig. 12A ) 1.00-1.10 times as long as broad; anterior margin between rostrum and lateral projections concave; anterolateral margins sloping or slightly terraced; posterior margin truncate; dorsal surface with four to six pairs of tufts of setae; paragastric grooves inconspicuous. Rostrum broadly triangular, rounded or terminating in acute or subacute spine, reaching or slightly overreaching lateral projections. Lateral projections obtusely triangular, with marginal or submarginal spinule. Posterior carapace similar to that of P. similis .

Ocular peduncle ( Fig. 12A ) 0.50-0.60 time as long as shield, weakly inflated basally; cornea not dilated, its maximum diameter 0.25-0.30 of length of ocular peduncle and subequal to basal diameter. Ocular acicle ( Fig. 12A ) narrowly triangular, slightly curved ventrally, terminating subacutely or bluntly and usually with slender submarginal spine.

Antennular peduncle ( Fig. 12A ) overreaching cornea by 0.30-0.50 length of ultimate segment. Ultimate segment 1.20-1.40 times longer than penultimate segment, slightly broadened distally in lateral view.

Antennal peduncle ( Fig. 7A ) overreaching distal margin of cornea by 0.20-0.40 length of fifth segment. Second segment with dorsolateral distal angle strongly produced, reaching midlength to distal margin of fourth segment, terminating in simple or bifid spine; dorsomesial distal angle with small spine. First segment laterally with small submarginal spine, ventromesial distal margin with few spinules laterally. Antennal acicle reaching or slightly overreaching distal margin of cornea, arcuate, terminating in acute spine; mesial margin with row of tufts of long stiff setae. Antennal flagellum longer than fully extended right cheliped, every article with some minute setae.

Mouthparts generally similar to those of P. similis . Third maxilliped with three to five (rarely two) accessory teeth on ischium ( Fig. 12B ).

Chelipeds grossly unequal. Right cheliped ( Fig. 13A ) with chela 1.40-1.70 times longer than greatest width at base of dactylus in females and small males, but noticeably elongate in large males, length attaining twice maximum width. Dactylus with closely-spaced, broad spines or tubercles on dorsal surface, showing imbricate appearance (some tubercles in proximal half of dactylus capsulate); dorsomesial margin nearly straight in lateral view, with row of moderately large, forwardly directed tubercles or spines; mesial face with numerous, low, broad spines; ventral surface with several low, squamiform tubercles and scattered tufts of moderately long stiff setae. Palm shorter than carpus; dorsomesial margin delimited by single or double row of moderately small spines; dorsolateral margin with row of small spines decreasing in size proximally; dorsal surface convex, with sparse tufts of long setae and numerous, small, capsulate tubercles and spinules, and also with row of moderately small spines on midline of dorsal surface extending onto fixed finger; corneous, spiniform capsules weakly curved backward, arising from anterior part of tubercles, basal pores rounded or “heart-shaped”; spinules and capsulate tubercles on dorsal surface of palm each with several short plumose setae arising from anterior bases; mesial face of palm flat or slightly concave, with scatter- ed low, squamiform protuberances; ventral surface with several low, broad protuberances. Carpus subequal in length to merus; dorsomesial margin distinctly delimited by row of moderately large spines; dorsal surface with numerous spinulose or capsulate tubercles and several moderately large spines adjacent to dorsolateral margin and with dense covering of short plumose setae; dorsolateral margin not delimited; lateral face with low, broad, sometimes multidenticulate protuberances dorsally, and small, low protuberances ventrally, ventrolateral distal margin smooth; mesial face slightly concave, ventromesial distal margin without row of spines; ventral surface with some spinulose tubercles. Merus with short transverse rows of setae on dorsal surface; dorsodistal margin lacking spine; mesial face somewhat inflated ventrally in large males, not inflated in small males and females; in large males, ventromesial margin produced, unarmed or armed only with few small spines proximally, and with numerous long setae; in females and small males, ventromesial margin not produced, with row of small spines and sparse setae; ventrolateral margin with row of small spines and numerous setae; ventral surface with scattered long setae, more strongly concave in large males than in females and small males. Ischium with smooth ventromesial margin and ventral surface smooth. Coxa without spines on distal margin.

Left cheliped ( Fig. 13B ) weakly compressed laterally. Chela elongate subovate in dorsal view, 2.80-3.20 times longer than greatest width at base of dactylus. Dactylus much longer than palm. Palm about half length of carpus, triangular in cross section; dorsal surface elevated in midline but not forming distinct ridge or crest, with row of moderately large spines extending onto proximal 0.20-0.40 of fixed finger; dorsolateral margin with row of small spines; dorsolateral and dorsomesial surfaces strongly sloping ventrally, former surface with covering of capsulate tubercles extending to proximal half of fixed finger, and latter surface with some capsulate tubercles dorsally and also with few small spinulose tubercles; dorsomesial margin not distinctly delimited. Carpus subequal in length to merus; dorsomesial margin with row of slender spines; dorsolateral margin weakly delimited, with row of four to six small spines; dorsal surface somewhat sloping, with several capsulate tubercles and tiny spinulose tubercles; dorsodistal margin with row of tiny spines; mesial face with scattered low protuberances, distomesial margin smooth; lateral face with numerous low, sometimes multidenticulate protuberances, ventrolateral distal margin not strongly expanded, distally with row of small spines or tubercles. Merus without spine on dorsodistal margin; ventromesial margin with few spinulose tubercles; lateral face ventrally with several small, low, somewhat squamous tubercles, ventrolateral margin not strongly expanded, with row of spines increasing in size distally; ventral surface weakly concave. Ischium and coxa similar to that of right cheliped.

Second and third pereopods ( Fig. 12 C-E) stouter than in P. similis . Dactyli 1.20-1.60 times as long as propodi; dactylus of right third pereopod 6.20- 7.77 times as long as high in males, 5.13-6.37 times as long in females; dorsal surfaces each with tufts of short setae and row of corneous spines increasing in length distally; ventral margins each with eight to 11 long corneous spines increasing in length distally. Propodi longer than carpi; dorsal surfaces each with short transverse ridges accompanied by rows of moderately short setae, but without spines. Carpi with two moderately small spines on dorsal surfaces of second, only with dorsodistal spine in third. Meri broad, ventrolateral distal margins unarmed.

Fourth pereopods with dactylus bearing row of fine corneous teeth on ventral margin; preungual process subequal in length to distalmost corneous spine, terminating bluntly, flexible. Propodal rasp composed of six or seven rows of corneous scales.

Males with four unpaired left (second to fifth) pleopods, all four unequally biramous. Females also with four unpaired left pleopods.

Telson ( Fig. 12F ) wider than long, with deep lateral indentations; posterior lobes, separated by small median cleft; terminal margins weakly oblique, each with row of five to nine closely spaced shot to long spines and several smaller submarginal spines; lateral margins smooth or weakly dentate.

COLORATION

In life: generally red or purplish red. Shield with patches of dark red laterally. Ocular peduncle generally white, with red band medially and with red patch at base of cornea. Antennular peduncle generally red; ultimate segment banded with white distally and proximally; penultimate segment white distally. Antennal peduncle generally red; fifth segment white dorsally and ventrally. Right cheliped with dark red or purple spot on mesial face centrally; small spines on dorsal surface of palm reddish; spines, spinules and tubercles on dorsal surface of carpus dark red; mesial surface of carpi with tinge of dark red proximally; lateral surface of merus with L-shaped line on lateral surface distally. Color and markings of left cheliped similar to that of right, but lacking dark red spot on mesial face of palm. Dactyli of ambulatory pereopods generally dark red, each with white patch slightly distal to midlength and red median stripe on lateral and mesial faces; propodi each with two dark red blotches, one on proximal half of dorsal surface and one at about midlength of ventral surface; lateral face with very short, faint, red longitudinal stripe distally, short obliquely transverse ridges (accompanied with setae) not or lightly colored with red; carpi each with large red blotches on lateral and ventral surface respectively; meri each with longitudinal white stripe and tinge of dark red dorsally.

SIZE

Males sl 10.4-21.0 mm; females sl 6.7-17.6 mm; ovigerous females sl 10.4-17.6 mm.

VARIATION

In both P. similis and P. rubrior n. sp., the right cheliped exhibits considerable variation in males, similar to that observed in P. japonicus . It shows a tendency of elongation, particularly in the chela, with increase of overall body size. In large males, the ventromesial margin of the merus is thicken- ed and sometimes noticeably produced ventrally, bearing numerous long setae. In females and small males, the ventromesial margins is not expanded or thickened, with sparse setae and a row of spines.

The dactylus of the right third pereopod tends to be more elongate in males than in females in both species ( Fig. 14 ). In P. similis , the proportional ratio “length/proximal depth” ranges from 8.00 to 10.30 (9.26 on average, N = 15) in males and from 7.20 to 9.16 (8.18 on average, N = 8) in females; in P. rubrior n. sp., the ratio ranges from 6.20 to 7.77 (6.59 on average, N = 21) in males and from 5.13 to 6.37 (6.09 on average, N = 14) in females. The difference in the mean values between male and female is significant in each species (in P. similis : t = 3.7026, d.f. = 20, p <0.01; in P. rubrior n. sp.: t = 2.6912, d.f. = 34, p <0.02).

REMARKS

Ortmann (1892) briefly described Eupagurus similis from a single male specimen collected in Kagoshima Bay, Kyushu, southern Japan, but did not illustrate his specimen. He compared this species with E. japonicus sensu Ortmann (1892) and separated the taxa by lengths of the antennal peduncle and antennal acicle as opposed to the ocular peduncle (longer in P. similis than in P. japonicus sensu Ortmann ), acuteness of the ocular acicle (more acute in the former than in the latter), development of a median row of spines on the right palm (less distinct in the former than in the latter), shape of the right palm (more elongate in the former than in the latter) and shape of the ambulatory dactyli (more elongate and slender in the former than in the latter). It has been found that most of these characters do not provide taxonomic significance in discriminating the two morphs of Pagurus similis s.l., because of variation. Nevertheless, the shape of the ambulatory dactyli are reliable in distinguishing the two morphs (see below). The dactyli are more elongate and slender in the orange morph than in the red morph. Therefore, the orange morph is considered to represent the true P. similis . The red morph, corresponding to P. japonicus sensu Ortmann , is described as new, P. rubrior n. sp.

Pagurus similis and P. rubrior n. sp. are very similar to each other. The most useful is the shape of the dactylus of the right third pereopod. The dactylus of the right third pereopod is much more slender in P. similis than in P. rubrior n. sp. ( Fig. 14 ). In males, the proportional ratio “length/proximal depth” ranges from 8.00 to 10.30 (9.26 on average, N = 15) in P. similis , from 6.20 to 7.77 (6.59 on average, N = 21) in P. rubrior n. sp.; in females, from 7.20 to 9.16 (8.18 on average, N = 8) in P. similis , from 5.13 to 6.37 (6.09 on average, N = 14) in P. rubrior n. sp. ( Fig. 14 ). Further, the ocular peduncle is proportionally longer in P. similis than in P. rubrior n. sp. (0.60-0.70 time as long as the shield versus 0.50-0.60 time as long). The tubercles on the palm of the right chela are much more numerous and denser in P. rubrior n. sp. than in P. similis (cf. Figs 13A ; 9A ). The number of the accessory teeth on the third maxilliped is generally greater in P. rubrior n. sp. than in P. similis , though it partially overlaps ( Fig. 15 ). The ischium of the third maxilliped is provided with three to six (most frequently three or four) accessory teeth on either side in P. rubrior n. sp., rather than one to three (most frequently one) in P. similis . The general color is bright yellow or light tan in P. similis , red or purplish-red in P. rubrior n. sp.; the short ridges accompanied with row of setae on the dorsolateral surfaces of the ambulatory propodi are dark red in P. similis , but not darkly colored in P. rubrior n. sp. In addition, Pagurus similis is known in deeper waters than P. rubrior n. sp. The present specimens of P. similis were collected from sublittoral depths ranging from 20 to 200 m, while the specimens of P. rubrior n. sp. were collected at depths of 5 to 30 m.

Pagurus hirtimanus is immediately distinguished from both P. similis and P. rubrior n. sp. by the strongly dilated corneas and the roundly convex dorsal surface of the left palm. In P. similis and P. rubrior n. sp., the dorsal surface of the left palm is strongly elevated in the midline. Further, the structure of the capsulate tubercles on the chela is different between P. hirtimanus , and the latter two species. In P. hirtimanus , the slender capsules arise from the central part of the tubercle, rather than the anterior slope of the tubercles in the latter two species. Pagurus capsularis differs from P. similis and P. rubrior n. sp. in the more dilated corneas, lack of dense dorsal covering of setae on the chelae that is seen in the latter two species, and the more strongly oblique terminal margins of the telson. As mentioned previously, P. pergranulatus is characteristic in the morphology of the right cheliped (see Remarks under P. japonicus ).

Doflein (1902) reported Eupagurus similis from Yokohama in Tokyo Bay. The two females used by Doflein (1902) (ZSM 303/1; size not measured) were reexamined, and found that they actually represent P. dubius ( Ortmann, 1892) .

Terao (1913) placed Eupagurus similis in the synonymy of E. japonicus , but he did not comment further. However, Asian workers have accepted that P. similis and P. japonicus are distinct. A part of the specimens identified as P. similis by Miyake (1978) and Miyake & Imafuku (1980) have been reexamined. As expected, P. similis and P. rubrior n. sp. are mixed in the material studied by Miyake (1978). The following three specimens represent the true P. similis : 1 ovig. (NSMT-CrR 1619, Miyake det. No. 294), 1 ovig. (NSMT-CrR 1757, Miyake det. No. 373) and 1 (NSMT- CrR 2220, Miyake det. No. 545); the other two specimens are P. rubrior n. sp.: 1 (NSMT-CrR 1372, Miyake det. No. 191) and 1 (NSMT- CrR 2012, Miyake det. No. 463). The three specimens studied by Miyake & Imafuku (1980) (OMNH-Ar 1798, 1799, 1826) are P. rubrior n. sp. Most of the published color photographs referred to as P. similis ( Suzuki 1971: pl. 34, fig. 4; Miyake 1975: pl. 115, figs 6, 9; 1978: pl. 2, fig. 3; 1982: pl. 42, fig. 2; Takeda 1986: unnumbered fig.; 1994: 228, fig. 5; Asakura 1995: pl. 97, fig. 4; Kobayashi 2000: unnumbered fig.; Minemizu 2000: unnumbered fig.) all actually depict P. rubrior n. sp.; only Yu & Foo (1990) shows the true P. similis . In listings, such as those of Miyake (1960, 1975, 1982, 1991, 1998), Suzuki (1971), Takeda (1982), which contain brief species accounts written in Japanese, my inclusion of the author’s citation as “in part” has been based on the species bathymetric range indicated. References by Alcock (1905) and Gordan (1956) are bibliographic treatments.

Kim (1964, 1970, 1973, 1985) reported P. similis from various locations in Korea. The photographed specimen ( Kim 1973: pl. 7, fig. 39) seems to represent P. rubrior n. sp. because of the relatively stout ambulatory legs. However, neither sufficient diagnostic information nor depth records were given in Kim’s accounts, and thus is difficult to know whether his material contained either P. similis , P. rubrior n. sp., or both, without direct examination of the material. M.-H. Kim and J.-N. Kim of Pukyon National University, Pusan, kindly provided me with a photograph depicting a specimen of Pagurus similis s.s. from Korean waters. The underwater photograph taken in Korean waters and published by Park & Choi (2001) clearly shows a specimen of P. rubrior n. sp. Therefore, there is no doubt that both species occur in Korean waters. Therefore, Kim’s references are questionably referred to P. similis .

Figures

FIG. 6. — A, Pagurus similis (Ortmann, 1892), sl 11.9 mm, from off Takeoka, Boso Peninsula (CBM-ZC 1058), entire animal in dorsal view; B, Pagurus rubrior n. sp., paratype sl 14.9 mm, from Takeoka, Boso Peninsula (MNHN-Pg 6099), entire animal in dorsal view.

FIG. 12. — Pagurus rubrior n. sp., holotype sl 13.2 mm, from off Hota, Boso Peninsula (CBM-ZC 4778); A, shield and cephalic appendages, dorsal (setae omitted from right); B, ischium of third maxilliped, dorsal (setae omitted); C, right second pereopod, lateral; D, dactylus and propodus of right third pereopod, lateral (setae omitted); E, left third pereopod, lateral; F, telson, dorsal (setae omitted). Scale bars: A, C-E, 5 mm; B, F, 2 mm.

FIG. 13. — Pagurus rubrior n. sp., holotype sl 13.2 mm, from off Hota, Boso Peninsula (CBM-ZC 4778); A, chela and carpus of right cheliped, dorsal (setae omitted); B, chela and carpus of left cheliped, dorsal (setae omitted). Scale bar: 5 mm.

FIG. 7. — Pagurus similis (Ortmann, 1892), sl 9.7 mm, from off Katsuyama Ukishima Islet, Boso Peninsula (CBM-ZC 3958); A, shield and cephalic appendages, dorsal (setae partially omitted from left; left antennular peduncle mutilated); B, carapace, dorsal (setae omitted from left); C, left ocular acicle, dorsal; D, left fourth pereopod, lateral; E, same, distal part of dactylus, lateral (setae omitted); F, coxae of fifth pereopods and eighth thoracic sternite, ventral; G, anterior lobe of sixth thoracic sternite, ventral; H, telson, dorsal. Scale bars: A, D, F, H, 2 mm; B, 5 mm; C, E, G, 1 mm.

FIG. 14. — Scatter plot showing proportional length of dactylus of right third pereopod (DL/DD) against shield length (sl) of Pagurus similis (Ortmann, 1892) and P. rubrior n. sp. Abbreviations: DL, length of dactylus; DD, proximal depth of dactylus.

FIG. 9. — Pagurus similis (Ortmann, 1892), sl 9.7 mm, from off Katsuyama Ukishima Islet, Boso Peninsula (CBM-ZC 3958); A, chela and carpus of right cheliped, dorsal (setae omitted); B, spines and tubercles on dorsal surface of palm of right chela, dorsal; C, capsulate tubercles on dorsal surface of right chela, mesial; D, same, dorsal; E, chela and carpus of left cheliped, dorsal (setae omitted). Scale bars: A, E, 5 mm; B, 1 mm; C, D, 0.5 mm.

FIG. 15. — Frequency distribution of number of accessory teeth on ischia of third maxillipeds of Pagurus similis (Ortmann, 1892) and P. rubrior n. sp. The numbers of the accessory teeth are indicated by combination of numbers counted from both sides. In the case of asymmetrical number of teeth, the arrangement does not precisely represent the actual side of the appendage, but only combination of numbers.