29 December 2000 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 113(4):945-955. 2000. Calappa pokipoki, a new species of box crab (Crustacea: Decapoda: Brachyura: Calappidae) from Hawaii

Peter K. L. Ng Department of Biological Sciences, National University of Singapore, Kent Ridge, Singapore 119260, Republic of Singapore

Abstract.—A new species of box crab, Calappa pokipoki, is described from Hawaii. This new species resembles C. gallus (Herbst), but can easily be dis­ tinguished by several carapace features as well as color pattern. Calappa po­ kipoki also resembles C. bicornis (Miers), C. matsuzawa Galil, C sebastieni Galil and C. yamasitae Sakai, but can be separated from these by a suite of distinctive carapace characters.

At present, seven species of calappids are Taxonomic Account known from Hawaii, Calappa calappa (Linnaeus 1758), C. gallus (Herbst 1803), Family Calappidae C. hepatica (Linnaeus 1758), C biconis Calappa pokipoki, new species (Miers 1884), Cycloes marierubri Galil & (Figs. 1, 2, 3a, 4a, 5a-g, 6) Clark, 1996, Mursia hawaiiensis Rathbun, Calappa bicornis.—Tinker, 1965: 74 (not 1893, and M. spinimanus Rathbun, 1906 Calappa bicornis Miers, 1884) (Rathbun 1906, Tinker 1965, Galil 1993, 1997; Galil & Clark 1996). Studies of the Material examined.—Holotype female brachyuran collections in the Barrice P. (87.8 by 64.5 mm) (BPBM 12073), Kona, Bishop Museum showed that Tinker's Hawaii, coll. research vessel Townsend (1965) record of C. bicornis is erroneous, Cromwell, TC 81-01 station 34, 16 Feb and the specimens which have been re­ 1981. ferred to this species actually represent a Paratypes: 1 female (78.8 by 58.8 mm) new species, described herein. This new (ZRC 2000.516), same data as holotype; 1 species, while most similar to C. gallus, dif­ male (43.5 by 33.5 mm) (BPBM SI 1259), fers from this species in several distinct TC 40 station 2, Penguin Banks, Molo- characters. kai, 21o09.9'-21°09.8'N, 157°24.1'- Specimens examined during this study 157°29.8'W, Hawaii, 182 m, coll. R/V are deposited in the Bernice P. Bishop Mu­ Townsend Cromwell, trawl, 6-7 Nov 1968; seum, Honolulu, Hawaii (BPBM); Zoolo- 1 juvenile (25.3 by 20.0 mm) (BPBM gisk Museum, University of Copenhagen SI 1255), TC 36 station 33, Penguin Banks, (ZMUC); and the Zoological Reference Molokai, 21°02.9'-21°01.6'N, 157°25.9'- Collection of the Raffles Museum, National 157°26.0'W, Hawaii, coll. R/V Townsend University of Singapore (ZRC). Numerous Cromwell, trawl, 5 Jun 1968; 1 female specimens of C. gallus, C. bicornis and C. (46.4 by 36.2 mm) (BPBM SI 1271), Bar­ galloides from BPBM, ZMUC and ZRC ber's Point, Oahu, 30 m, Hawaii, coll. 1963; have also been examined for the present 1 juvenile (13.3 by 11.0 mm) (BPBM comparisons. Measurements provided are SI 1146), Northwest End, Penguin Banks, listed as the carapace width by length. The Molokai, 21°02.8'N, 157°40.5'W, 182 m, abbreviations Gl and G2 are used for the Hawaii, coll. HURL 82-111, Makalii sub­ male first and second pleopods respectively. mersible, 20 Sep 1982; 1 male (42.4 by 946 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

31.6 mm) (ZRC 2000.1798), Molokai, Ha­ prominently swollen; outer part with 2 deep waii, coll. 14 Jun 1959; 1 female (right part narrow fissures. Eyes folding obliquely. Ex­ of carapace damaged) (BPBM S6688), Ha­ ternal orbital tooth very low, rounded. An­ waii, no other data; 1 male (49.6 by 36.4 terolateral margin arcuate, clearly demar­ mm) (BPBM S5422), 400 yards off Mok- cated from posterolateral margin by distinct ulua, Oahu, 36 m, Hawaii, coll. Brock, 14 cleft, lined with scattered long, stiff yellow Apr 1949. setae which partially obscure margin; first Diagnosis.—Carapace dorsal surface tooth (subhepatic tooth) large, low, conical; covered with relatively large, low, rounded, subsequent 5 teeth small but clearly dis­ submammillate granules on anterior half; cernible, with bicuspid tips, lateral margins posterior half with numerous oblique to with minute denticles; last 6 teeth ca. twice transverse raised strise; hepatic region as large as preceding teeth, with sharp tips, strongly depressed; from posterior view, lateral margins distinctly denticulate. Pos­ gastric region is highest point, with bran­ terolateral clypeiform extension well devel­ chial regions gradually sloping towards oped, lined with numerous long, stiff yel­ margins; frontal margin quadridentate; an­ low setae which almost completely obscure terolateral margin with 11 or 12 well margin; anterior part with 3 broadly trian­ marked teeth; clypeiform extension well de­ gular teeth (larger posteriorly), each tooth veloped, margin with numerous long, stiff with denticulate margins and median ridge; yellow setae which almost completely ob­ posterior part with 3 low, broadly triangular scure margin; basal antennal segment with teeth, first tooth longest, second tooth distal margin gently concave; outer surface slightly shorter than first, third tooth (near­ of manus with well spaced granules; Gl est posterior carapace margin) ca. 0.5 times with distal \b slightly sinuous; G2 subequal length of second tooth; each tooth with in length to Gl, distal segment relatively strongly denticulated margins and median short, tip of flange between distal and basal ridge, margin appearing prominently serrat­ segments faintly tricuspid. ed; margins lined with long, stiff yellow se­ Description.—Carapace subtriangular in tae which almost obscure margin. Posterior shape; dorsal surface distinctly convex; an­ carapace margin convex, margin distinctly terior half covered with relatively large, denticulated, appears serrated, lateral parts low, rounded, submammillate granules lined with long, stiff yellow setae. Basal an­ which are closely appressed basally; pos­ tennal segment subtriangular, surface finely terior half with numerous oblique to trans­ granulated, distal margin gently concave. verse raised striae which are longer towards Longitudinal endostomial septum with shal­ lateral margins; striae near margins more low concavity on anterior border, edge just prominent, beaded, appearing almost ser­ visible when first maxillipeds closed. rated. Hepatic region strongly depressed. Minor left chela (without cutting tooth) Epigastric region with 2 distinct transverse with prominent dorsal crest, with 6 lamel- tubercles. Gastric region strongly raised, liform teeth, first bilobed, low, next 5 high, median part especially high. Gastric and acutely triangular; outer surface of manus cardiac regions separated from branchial re­ granulated, granules not arranged in distinct gions by deep longitudinal groove. Subor­ rows; upper half with several large gran­ bital region with 2 large transverse gran­ ules; lower Vz surface with numerous subs- ules, rest of surface finely granulated. Fron­ quamose granules, becoming smaller to­ tal margin thickened, quadridentate, median wards tip of pollex; subsquamiform gran­ part prominently produced anteriorly, me­ ules on proximal lower surface of manus dian teeth separated by distinct U-shaped becoming larger and fewer towards carpus; sulcus. Supraorbital margin with small but ventral outer margin lined with row of lat­ distinct inner tooth; inner half of margin erally directed granules, appears distinctly VOLUME 113. NUMBER 4 947

Fig. 1. Calappa pokipoki, new species, holotype female (87.8 by 64.5 mm) (BPBM S5422). a, dorsal view b, posterior carapace margin (dorsal view): c, right chela (outer view). 948 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Calappa pokipoki, new species, paratype male (42.4 by 31.6 mm) (ZRC 2000.1798). a. dorsal view; b, posterior carapace margin (dorsal view); c, chelipeds (outer view). VOLUME 113. NUMBER 4 949

Fig. 3. Carapace showing degree of convexity (posterior view), a, Calappa pokipoki, new species, paratype male (42.4 by 31.6 mm) (ZRC 2000.1798): b. C. gallus, male (48.4 by 36.4 mm) (ZRC). serrated, tooth adjacent to carpus low but stiff setae; subproximal part of ventral mar­ distinct, subtruncate, sublamelliform; dac­ gin of merus with short row of long stiff tylus very slender, recurved, distal half pig­ setae. mented, smooth; proximal part of dactylus Abdomen narrowly triangular. First seg­ granulated. Major right chela (with special ment with proximal margin lined with sub- cutting tooth) similar in shape and features median low granules, appearing gently ser­ to minor chela, but dorsal crest, with 7 la- rated, with long, stiff yellow setae, outer­ melliform teeth, first bilobed, low, next 6 most parts smooth, glabrous. Second seg­ high, acutely triangular; entire dactylus pig­ ment with distinct granules on outer and mented, proximal surface granulated, with median parts, lined with long, stiff yellow subtruncate tooth proximally. setae, intermediate areas unarmed, gla­ Ambulatory legs relatively slender, un­ brous. Third to fifth segments fused, but lat­ armed, smooth; second pair longest. Dac­ eral clefts separating segments still discern­ tylus very slender, styliform, gently curved. ible; third segment subrectangular, outer Last leg with subdistal part of ventral mar­ submarginal part with scalloped features, gin of fused basis-ischium with tuft of long fourth segment trapezoidal, lateral margins 950 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Posterior carapace margin, first two male abdominal segments and base of last pair of ambulatory legs, a, Calappa pokipoki, new species, paratype male (42.4 by 31.6 mm) (ZRC 2000.1798); b, C. gallus, male (48.4 by 36.4 mm) (ZRC).

deeply concave; fifth segment almost squar­ in larger specimens, regardless of sex. One ish, with deeply concave lateral margins; paratype female (78.8 by 58.8 mm, ZRC sixth segment slightly elongate, lateral mar­ 2000.516) is very unusual in that the spe­ gin sinuous. Telson acutely triangular, lat­ cial cutting tooth is on the left chela rather eral margin gently concave to almost than the right. In all known specimens of straight. all other species of Calappa Weber, 1795, Gl relatively stout, almost straight, distal as well as all the other specimens of C. po­ V* slightly sinuous, more slender than me­ kipoki, the special cutting tooth is on the dian part; tip truncate; distal surfaces lined right chela and is adapted to peel right-aper- with numerous small posteriorly directed tured gastropods (see Ng & Tan 1984, spinules. G2 subequal in length to Gl, gent­ 1985). ly curved; distal segment relatively short, Color (in preservative).—In adult speci­ tip rounded; junction between basal and mens, anterior half of carapace orange; pos­ distal segments with short flange, tip of terior xh dirty white with scattered uneven flange faintly tricuspid; inner margin of brown spots; zone where 2 color zones base of basal segment bilobed. meet uneven but well marked. Despite the Variation.—The specimens do not vary age of the present specimens, most of them substantially in morphology, with the non­ still show the pattern described above quite sexual characters relatively constant. The clearly. setation on the carapace margin, ambula­ Etymology.—The species name is de­ tory legs and abdomen is less dense in rived from the Hawaiian name for Calappa, small specimens (less than 30 mm carapace "Poki-poki." The name is used here as a width). The chela is also relatively higher noun in apposition. VOLUME 113. NUMBER 4 951

Fig. 5. a-g, Calappa pokipoki, new species: a, male abdomen; b, left basal antennal segment; c, 1, left second ambulatory leg (dactylus and propodus); d-k, frontal margins; a, b, c, d, holotype female (87.8 by 64.5 mm) (BPBM S5422); e, male (43.5 by 33.5 mm) (BPBM SI 1259); f, juvenile (25.3 by 20.0 mm) (BPBM SI 1255); g, juvenile (13.3 by 11.0 mm) (BPBM SI 1146); h-1, C. gallus; h, female (47.2 by 36.2 mm) (BPBM 5078); i, female (46.2 by 35.8 mm) (BPBM 967); j. male (33.2 by 25.5 mm) (BPBM 3303); k, female (46.8 by 37.8 mm) (BPBM 1303). Scales: a, c, 1 = 5.0 mm; b, d-k = 1.0 mm.

Remarks.—Calappa pokipoki, new spe­ BPBM has a good series of specimens of cies, is most similar to C. gallus (Herbst C. gallus which allows for a very thorough 1803), a common species in Hawaii, but the competition of the two species to be made. two can easily be distinguished by the fron­ The frontal margin of C. pokipoki (irre­ tal margin and carapace physiognomy. The spective of sex or size) is more produced, 952 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Calappa pokipoki, new species, paratype male (49.6 by 36.4 mm) (BPBM S5422). a, left Gl (dorsal view); b, left Gl (ventral view) (setae not drawn); c, distal part of left Gl (dorsal view); d, distal part of left GI (ventral view); e, left G2 (ventral view); f. flange at junction of basal and distal segments of G2; g, G2 basal segment. Scales: a, b, e = 1.0 mm; c, d, f, g = 0.5 mm. with four well defined teeth, the two me­ species. Specimens of C. pokipoki larger dian ones projecting anteriorly from the lat­ than 40 mm carapace width can also easily eral ones and are separated by a distinct be distinguished from C. gallus by the cleft (Fig. 5d-g). In C. gallus, however, the much denser and longer setae lining the frontal margin is subquadrate, and although posterolateral margins and chelipedal meral the lateral parts may appear stepped, it nev­ lobe which almost completely obscure the er develops even a trace of a tooth (Fig. 5h- margins (Fig. 4a) from the dorsal view. In k). The carapace of C. pokipoki is distinctly C. gallus, these setae are much less dense less swollen compared to that of C. gallus. even in large specimens, and never com­ Observed frontally or from the posterior, pletely obscure the margins (Fig. 4b). The the branchial regions of C. pokipoki in par­ ambulatory dactyli of C. pokipoki is also ticular, are prominently lower and gradually relatively more slender and somewhat lon­ slope towards the lateral margins from the ger (Fig. 5c) compared to those of C. gallus gastric region which is the highest point of (Fig. 51). The second male abdominal seg­ the carapace (Fig. 3a). For C. gallus, the ment is also distinctly more granulose in C. gastric and branchial regions are equally gallus (Fig. 4b vs. 4a). In contrast, the pos­ high (Fig. 3b, see also Galil 1997, Sakai terolateral margin of C. pokipoki has more 1999). widely spaced granules (Figs. 3a, 4a vs. 3b, In addition, several more subtle charac­ 4b). In C. gallus, the setose parts of the ters can also be used to distinguish the two basis-ischium and merus of the last ambu- VOLUME 113, NUMBER 4 953 latory leg have gentle but distinctly scal­ form of the frontal margin, relatively more loped granules (Fig. 4b), but in C. pokipoki expanded clypeiform process, granulation these parts are almost smooth (Fig. 4a). features of the suiter surface of the chela There also appear to be color differences and structure of the Gl, C. pokipoki closely between C. pokipoki and C. gallus. Calap- resembles C. sebastieni Galil, 1997, known pa pokipoki clearly has a "bicolored" car­ only from the Marquesas. Calappa sebas­ apace when adult, with the anterior half of tieni can easily be separated from C. poki­ the carapace darker colored and the poste­ poki by its less broad carapace (length to rior part white with scattered brown spots, width ratio 1.6 vs. 1.3) and much smoother with the two zones more or less clearly de­ dorsal carapace surface which does not marcated. This color pattern is similar to have any prominent large granules on the that reported for species like C. undulata anterior part of the carapace (Galil 1997: Dai & Yang, 1991, and C. bicornis Miers, 312, fig. 17f). In the form of its dorsal car­ 1884 (see Miyake 1983, Chen 1993, Ng et apace sculpture, C. pokipoki superficially al. 2000). resembles C. yamasitae Sakai, 1980, known In the form of the frontal margin (and only from Japan, but the latter species can general color pattern), C. pokipoki certainly be separated by its proportionately broader resembles C. bicornis, and it is not surpris­ carapace (width to length ratio 1.6 vs. 1.3), ing that Tinker (1965) referred it to this spe­ bidentate (vs. quadridentate) frontal margin cies. Tinker's (1965) material is in the and the dorsal crest of larger chela having BPBM and his figure is probably based on more teeth (9-10 vs. 6-7) (cf. Sakai 1980: one of the specimens listed above. Calappa 5, frontispiece fig. 2). bicornis, however, can easily be distin­ Calappa pokipoki also bears a close re­ guished from C. pokipoki by the dorsal car­ semblance to the Atlantic species previous­ apace surface of C. bicornis having larger ly referred to "Calappa gallus" (see Mo- and/or more mammillate tubercles, less ex­ nod 1956, Manning & Holthuis 1981). panded clypeiform process, relatively much Manning & Chace (1990) referred the old deeper cleft on the frontal margin, deeply records of "Calappa gallus" (nee Herbst clefted anterior margin of the basal antennal 1803) from the Atlantic to C. galloides segment which appears bidentate (vs. gent­ Stimpson, 1859. I have examined speci­ ly concave), serrated supra- and suborbital mens from various parts of the Atlantic and margins (vs. smooth), proportionately more there appears to be at least two species re- slender ambulatory meri and much longer ferrable to "C. galloides" from there; one dactyli, with the legs appearing banded in with a prominently quadridentate frontal life (vs. uniformly colored and patterned) margin and less inflated carapace (C gal­ and the distal segment relatively more pod­ loides s. str.); and another with a more like (vs. evenly cylindrical with a rounded rounded frontal margin and strongly inflat­ tip) (present specimens, see also Galil 1997, ed carapace. In addition, there is one more Ng et al. 2000). supposed junior synonym of C. galloides, In the morphology and sculpture of the C. squamosa Desbonne in Desbonne & carapace, C. pokipoki also resembles C. Schramm, 1867, described from Guade­ matsuzawa Galil, 1997, described and loupe. Although the resolution of the tax­ known only from Japan thus far. The cly­ onomy of taxonomy of these Attantic taxa peiform process of C. matsuzawa, however, is well outside the scope of the present pa­ is relatively less expanded, the anterior part per, it is nevertheless useful to note that C. of the anterolateral margin has no well pokipoki is very unlikely to be conspecific marked teeth (vs. with 6 distinct teeth in C. with any of the above taxa. Calappa poki­ pokipoki), and the front is less well devel­ poki can be separated from C. galloides oped (Galil 1997:304, Fig. 17b). In the (and C. squamosa) by its relatively more 954 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON expanded clypeiform extension, more moires du Museum national d'Histoire Naturel­ rounded posterior carapace margin and le, Paris 176:271-335. , & P. F. Clark. 1996. A revision of Crypiosoma much higher dorsal crest of the chela. Brulle, 1837 and Cycloes de Haan, 1837 (Crus­ Calappa pokipoki is known only from tacea brachyura: Calappidae).—Zoological the Hawaiian islands and its vicinity thus Journal of the Linnaean Society 117:176-204. far. All the type specimens have been col­ Herbst, J. F. W. 1782-1804. Versuch einer Naturges- lected from sublittoral waters. Calappa gal- chichte der Krabben und Krebse, nebst einer systematischen Beschreibung ihrer verschiede- lus on the other hand, has a very wide Indo- nen Arten. vol. 1-3, 515 pp., 62 pis. West Pacific distribution and is common Linnaeus, C. 1758. Systema Naturae per regna tria na­ from intertidal to deeper waters. turae, secundum classes, ordines, genera, species, cum characteribus, differentiis specierum, syn- onymis, locis. 10th edition, l:i-iii, 1-824 pp. Acknowledgments Manning, R. B., & L. B. Holthuis. 1981. West African brachyuran crabs (Crustacea: Decapoda).—Smith­ I am most grateful to Lu Eldredge who sonian Contributions to Zoology 306:1-379. invited me to Hawaii to work on the crabs , & F. A. Chace, Jr. 1990. Decapod and sto- in the BPBM, and the Edmondron Fund for matopod Crustacea from Ascension Island, supporting the visit. The hospitality and South Atlantic Ocean.—Smithsonian Contribu­ tions to Zoology 503:1-91. kindness of all the folks in the BPBM also Miers, E. J. 1884. Crustacea. Pp. 1178-1322 in Report is very much appreciated. Torben Wolff and on the zoological collections made in the Indo- Danny Jacobsen (ZMUC) kindly allowed Pacihc Ocean during the voyage of H.M.S. me access to their valuable collections. 'Alert' 1881-2. British Museum, London. Miyake, S. 1983. Brachyura (Crabs). Japanese crusta­ cean decapods and stomatopods in Color, 2:277 Literature Cited pp., pis. 1-64, unnumbered fig. 1st edition. Hoikusha, Osaka. [In Japanese] Chen, H.-L. 1933. The Calappidae (Crustacea: Brach- Monod, T. 1956. Hippidea et Brachyura ouest-afri- yura) of Chinese waters. Pp. 675-704 in B. cains.—Memoires Institut franc,ais d'Africa Morton, ed.. Marine biology of the South China Noire 45:1674. Sea, Proceedings of the First International Con­ Ng, P. K. L., J. C. Y Lai, & C. Aungtonya. 2000. The ference on Marine Biology, Hong Kong, South box and moon crabs of Thailand, with descrip­ China Sea, Hong Kong University Press. tion of a new species of Culappa (Crustacea: Dai, A.-Y., & S.-L. Yang. 1991. Crabs of China Seas. Brachyura: Calappidae, Matutidae).—Research Pp. 21+682, figs. 1-295, pis. 1-74. China Bulletin of the Phuket Marine Biological Cen­ Ocean Press, Beijing & Springer-Verlag, Berlin. ter, Thailand, (in press). Desbonne I., & A. Schramm. 1867. Brachyures. In , & L. W. H. Tan. 1984. The 'shell peeling' Crustaces de la Guadeloupe, d'apres un man- structure of the box crab Calappa philargius uscrit du Doctaur Isis Desbonne compare avec (L.) and other crabs in relation to mollusc shell les echantillons de Crustaces de sa collection et architecture.—Journal of the Singapore Nation­ les dernires publications de MM. Henri de Saus- al Academy of Science 13:195-199. sure et William Stimpson, Ire partie, Basse-Ter- , & . 1985. "Right Handedness' in het- re. 60 pp. 8 pis. erochelous calappoid and xanthoid crabs—sug­ Fabricius, J. C. 1793. Entomologia systematica emen- gestion for a functional advantage.—Crusta- data et aucta secundum classes, ordines, genera, ceana 49:98-100. species adjectis synonimis, locis, observationi- Rathbun, M. J. 1893. Descriptions of new genera and bus descriptionibus. Hafniae 2:i-viii, 1-519. species of crabs from the West Coast of North Galil. B. 1993. Crustacea Decapoda: A revision of the America and Sandwich Islands. In Scientific genus Mursia Desmarest, 1823 (Calappidae). In Results of Exploration by the U.S. Fish Com­ A. Crosnier, ed., Resultats des Campagnes MU- mission 'Albatross.' No. XXIX.—Proceedings SORSTOM, vol. 10.—M6moires du Museum of the U.S. National Museum 16(933):223-260. national dHistoire Naturelle, Paris 156:347- . 1906. The Brachyura and Macrura of the Ha­ 379. waiian Islands.—Bulletin of the United States . 1997. Crustacea Decapoda: A revision of the Fish Commission 23:827-930, pis. 1-24. Indo-Pacihc species of genus Calappa Weber, Sakai, K. 1999. J. F. W. Herbst-Collection of decapod 1795 (Calappidae) In A. Crosnier, ed., Resultats Crustacea of the Berlin Zoological Museum, des Campagnes MUSORSTOM, vol. 18.—Me- with remarks on certain species.—Naturalists, VOLUME 113, NUMBER 4 955

Tokushima Biological Laboratory 6:1-45, pis. Tinker, S. W. 1965. Pacific Crustacea. An illustrated 1-21. handbook on the reef-dwelling Crustacea of Ha- > ? Sakai, T 1980. New species of crabs of the families waii and the South Seas. C. E. Tuttle, Tokyo Lithodidae and Calappidae.—Researches on 134 pp., 52 pis. Crustacea, Tokyo 10:1-11, frontispiece, pi. 1. Weber, E 1795. Nomenciator entomologicus secundum Stimpson, W. 1959. Notes on North American Crus­ Entomologiam systematicam ill. Fabrict adjects tacea, No. 1.—Annals of the Lyceum of Natural speciebus recens detectis et varietatibus, i—viii, History of New York 7:49-93, pi. 1. 1-171.