Aparatanais hawaiensis, Morales-Núñez, Andrés G., Pelleteri, Sara & Heard, Richard W., 2016

Morales-Núñez, Andrés G., Pelleteri, Sara & Heard, Richard W., 2016, Two new paratanaid Tanaidacea (Crustacea: Malacostraca: Peracarida) from the Hawaiian Islands, with illustrated taxonomic keys, Zootaxa 4150 (4), pp. 467-492 : 477-489

Aparatanais hawaiensis sp. nov.

Figs. 8−16 , 17 B, 18D

Material examined. Holotype— non-ovigerous ♀, length 3.9 mm, ( BPBM S16459 ), Station (Stn) 11(21º 16' 26.5"N; 157º 50' 26.1"W), Kaizers Channel, Waikiki , O'ahu, Hawaii, depth 10−12 m, 22 January 2001.

Paratypes— two ♀♀ ( BPBM S16460 ), same collection data as for holotype ; two ♀♀ ( USNM 1411098 ), Stn 10 (21º 15' 34.8"N; 157º 49' 39.7''W), Kapua Channel, Waikiki , O'ahu, Hawaii, depth 0−5 m, 22 January 2001: one ♂ ( BPBM S16460 ), same collection data as for holotype. Additional specimens from the type locality are in the collection of the authors.

Additional material— one ♀, Stn 1, 21º 30' 22.1"N; 157º 50' 57.1"W, KYC Patch Reef, Kaneohe Bay , O'ahu, Hawaii, depth 4−6 m, 11 May 1999 ; two ♀♀, Stn 3 (21º 30' 3.9"N; 157º 50 25.7"W), Hakip'ui Reef, Kaneohe Bay , O'ahu, Hawaii, depth 1.5−5 m, 17 November 17 1999 ; one ♂ and one ♀, Stn 9 (21º 27' 35.2"N; 157º 49' 22.1"W), Channel Marker 12 Reef , Kaneohe Bay , O'ahu, Hawaii, depth 5−15 m, 19 January 1999 ; eight ♀♀, Stn 25, 21º 28'0"N; 157º 50'0"W, Moku Manu Island, Kaneohe Bay , O'ahu, Hawaii, depth 18 m, 26 January 2000 ; four neuters, Stn 1 (21º 16' 35.5'N; 157º 50' 26.4"W), Kaizer's Wreck, Waikiki , O'ahu, Hawaii, depth 1.5−5.5 m, 15 January 2001 ; ten ♀♀, Stn 2, 21º 16' 10.8"N; 157º 50' 1.3"W, Outside Pops, Waikiki , O'ahu, Hawaii, depth 6 m, 15 January 2001 ; four ♀♀, Stn (21º 15' 17.4"N; 157º 49' 19"W), Toggs Wreck, Waikiki , O'ahu, Hawaii, depth 4 m, 18 January 2001 ; three ♀♀ and six neuters, Stn 5 (2º 15'53.5"N; 157º 49' 36"W), Aquarium outside reef, Waikiki , O'ahu, Hawaii, depth 3 m, 17January 2001 ; four neuters, Stn 6 (21º 15' 37.6"N; 157º 50'15.3"W), Atlantis Wreck, Waikiki , O'ahu, Hawaii, depth 30−40 m, 18 January 18 2001 ; one ♂, one ♀ and five neuters, Stn 10 (21º 15' 34.8"N; 157º 49' 39.7''W), Kapua Channel, Waikiki , O'ahu, Hawaii, depth 0−5 m, 22 January 2001 ; nine subadult female and eight neuters, Stn 11 (21º 16' 26.5"N; 157º 50' 26.1"W), Kaizers Channel, Waikiki , O'ahu, Hawaii, depth 10−12 m, 22 January 2001 ; seven ♀♀, Stn 15 (21º 15' 34.8 N; 157º 49' 39.7" W), Ft. DeRussy Reef, Waikiki , O'ahu, Hawaii, depth 1.5 m, 24 January 2001 .

Diagnosis. Female: Body elongate, about eight times as long as wide. Carapace superficially subdivided “plates.” Pleonites 1-4 with pair of proximally swollen setulose setae on ventrolateral margins. Antenna article-2 with dorsal margin having dense row of minute, hirsute setae. Maxillule with endite terminating in eight spiniform setae. Maxilliped palp article-2 with inner margin armed with narrow, blade-like spiniform seta bearing four, low, broadly spaced denticles on ventral margin. Cheliped inner face of propodus having one short, stout, subdistal, setulate setae adjacent to articulation with dactylus.

Male: Antennule with seven articles, three peduncular and four flagella. Cheliped palm with inner face having “comb row” of ten stout setae just proximal to articulation with dactylus. Uropod endopod article-1 having a row of transverse sensory-setae on inner mid-margin, and article-2 longer than endopod article-1.

Etymology. The species is named after the Hawaiian Islands where the specimens used in this study were collected.

Type locality. Kaizers Channel, Waikiki, (21º 16' 26.5"N; 157º 50' 26.1"W), O'ahu, Hawaii.

Distribution. Presently known with certainty from the Hawaiian island of O’ahu.

Description. Non-ovigerous female.

Body ( Figs. 8 A −B, 16 A −B): Length about 3.5 mm, about eight times width.

Cephalothorax ( Figs. 8 A −B, 16 A −B): About 14% of TL, with length shorter than pereonites 1−2 combined, with simple seta near each ocular lobe; demarked lines or sutures on carapace indicating possible union of lateral plates. Ocular lobes bearing pigmented eyes.

Pereon ( Figs. 8 A, 16 A −B): About 60% of TL, pereonite-1 shorter than other pereonites; pereonites 4 and 5 longest; all pereonites wider than long, with simple seta on each dorsal lateral margin.

Pleon ( Figs. 8 A, 16 A −B): About 18% of TL, shorter than pereonites 5‒6 combined; pleonite-1 slightly longer than the other pleonites; pleonites 2−5 subequal; pleonites 1−4 each with pair of swollen lateral, setulose setae ( Figs. 8 B −C, 16B) and pairs of short simple setae ( Fig. 8 A); pleonites-5 with lateral pair of short simple setae ( Fig. 8 B).

Pleotelson ( Figs. 8 A, 16 A −B): About 8% of TL, longer than pleonite-5, and distally with two sensory-setae and two simple setae.

Antennule ( Fig. 9 A): Shorter than carapace. Article-1 about 1.9 times as long as width; with two clusters of three and two sensory-setae (one medial and one sub-distal) and medial simple seta on dorsal margin. Article-2 slightly longer than wide, shorter than half-length of article-1, with two simple setae on sub-distal dorsal margin; ventral margin with sub-distal lateral seta. Article-3 about 1.8 times as wide as long, with one dorsal and one ventral simple setae of varying length on sub-distal margin. Article-4 elongate, length about 2/3 of article-1, with simple seta on distal inner margin. Small terminal, cap-like article with one sensory-seta, one aesthetasc, and two simple setae of varying lengths.

Antenna ( Fig. 9 B): Article-1 reduced, asetose. Article-2 about 1.4 times as long as wide, dorsal margin with dense row of minute setae terminating distally in long spiniform seta; mid-distal lateral margin with small seta; ventral margin forming shallow medial apophysis with weakly buttressed straight seta. Article-3 sub-quadrate, about 1.2 times as wide as long, with stout, distodorsal process. Article-4 about 2.5 times as long as wide, with one sensory-seta on mid-lateral margin, two sensory-setae and two small spiniform setae on sub-distal lateral margin. Article-5 about 2.2 times as long as wide, with one sub-distal lateral simple seta. Article-6 minute, with four simple setae of varying lengths.

Mouthparts: Labrum ( Fig. 9 C): Hood-shaped, distal edge finely setose. Mandibles ( Fig. 9 D −F): Molar process well developed; left mandible with smooth, strong incisor without crenulated upper margin, lacinia mobilis with four shallow denticles on ventral margin and cup-shape tip ( Fig. 9 E); right mandible with strong incisor with crenulate upper margin and bifid tip ( Fig. 8 F). Labium ( Fig. 9 G): Bilobate, finely setose distal margin. Maxillule ( Fig. 9 H): Endite with eight distal spiniform setae, outer and inner margin with short simple setae; palp slender with two long terminal setae of unequal length. Maxilla ( Fig. 9 I): Oval.

Maxilliped ( Fig. 10 A −B): Basis fused; long simple seta near articulation with palp, extending distally to or slightly beyond palp article-2. Endites broad, fused proximally, inner distal margin with simple seta, microtrichia and two small tubercles rounded; outer and medial distal margins serrate. Palp: article-1 asetose; article-2 triangular with small simple seta on outer proximal margin, inner margin with two finely pectinate (one long, one short) spiniform setae and modified spiniform seta with about four denticles on ventral margin, with cluster of setules on dorsal margin ( Fig. 10 B); article-3 with four setulate (one strongly curved) setae ( Fig. 10 A); article-4 with four setulate setae on inner distal margin, simple seta on outer subdistal margin, and cluster of minute setae on inner margin ( Fig. 10 A). Epignath: Not recovered.

Cheliped ( Fig. 10 C −F): Coxal sclerite sub-triangular, dorsally inserted. Basis about 2.8 times as long as wide, with simple seta on outer distodorsal margin. Merus triangular, with simple seta on mid-ventral margin. Carpus about 1.6 times as long as wide, with two (one sub-proximal and one sub-distal) short simple setae on dorsal margin, and two simple setae on sub-distal, ventral margin. Propodus about 1.2 times as long as width, palm about twice as long as fixed finger, with short setulate setae on medial inner margin near articulation with dactylus ( Fig. 10 E‒F); fixed finger with two ventral setae and three simple setae on outer incisive margin; broad kukri-shaped sinuate spiniform seta on medial margin near articulation with dactylus ( Fig. 10 D). Dactylus slightly longer than fixed finger, curved distally, unguis not fused; with simple seta proximally on inner sub-dorsal margin.

Pereopod-1 ( Fig. 11 A, G): Longer than other five pereopods. Coxa with simple seta. Basis slender, 3.4 times as long as wide, with dorsoproximal simple seta. Ischium wider than long with single simple ventral seta. Merus about 3.2 times as long as wide, with two ventrodistal small simple setae. Carpus about 2.4 times as long as wide, with three (two dorsal and one ventral) simple setae of unequal length on sub-distal margin. Propodus about 4.8 times as long as wide, with two sub-distal pectinate spiniform setae on dorsal margin ( Fig. 11 G) and simple seta on sub-distal ventral margin. Dactylus and unguis combined slightly shorter than propodus, dactylus with simple proximal seta, unguis longer than dactylus.

Pereopod-2 ( Fig. 11 B): Coxa with simple seta. Basis stout, 2.5 times as long as wide, with dorsoproximal simple seta. Ischium wider than long, with single simple ventral seta. Merus about 1.8 times as long as wide, with simple seta and one spiniform seta, distoventrally. Carpus about 1.7 times as long as wide, with small seta on subdistal dorsal margin and one longer distodorsal spiniform seta, with three distoventral spiniform setae of unequal length. Propodus about 3.6 times as long as wide, with one pectinate spiniform seta and small simple seta on subdistal dorsal margin and one sub-distal simple seta ventrally. Dactylus and unguis combined slightly longer than propodus and not fused, dactylus with simple proximal seta, unguis longer than dactylus.

Pereopod-3 ( Fig. 11 C, H‒J): Shorter than pereopods 1 and 2. Coxa with simple seta. Basis stout, 2.3 times as long as wide, with dorsoproximal simple seta. Ischium wider than long with single simple ventral seta. Merus about 2.7 times as long as wide, with simple seta and one spiniform seta, distoventrally. Carpus about 1.5 times as long as wide, with small seta on sub-distal dorsal margin and one longer distodorsal spiniform seta ( Fig. 11 H), with two distoventral spiniform setae of unequal length ( Fig. 11 I −J, respectively). Propodus about 2.9 times as long as wide, dorsal margin with microtrichia, with one small simple seta and pectinate spiniform seta on sub-distal dorsal margin, and one sub-distal simple seta ventrally. Dactylus and unguis combined approximately same length of that propodus and not fused, dactylus with simple proximal seta, unguis longer than dactylus

Pereopod-4 ( Fig. 11 D, K‒M): Basis stout, 1.9 times as long as wide, with simple seta and two sensory-setae on sub-distal ventral margin ( Fig. 11 K). Ischium wider than long, with two ventral simple setae. Merus length about twice as long as wide, with two short asymmetrical bifurcate spiniform setae on distoventral margin ( Fig. 11 L), and row of setules on half of ventrodistal margin. Carpus about 1.3 times as long as wide, with simple distodorsal seta ( Fig. 11 M), and four stout bifurcate modified spiniform setae, and row of setules on half of ventrodistal margin. Propodus about 3.1 times as long as width, with mid-dorsal sensory-seta and dorsodistal pectinate seta, with two bifurcate spiniform setae on ventrodistal margin, and row of setules on half of ventrodistal margin. Dactylus and unguis claw-like, together almost half-length of propodus, dactylus longer than unguis, curved and not fused, dactylus with row of setules on half of ventrodistal margin.

Pereopod-5 ( Fig. 11 E): Similar to pereopod-4, except basis and propodus slightly longer. Carpus slightly shorter. Propodus with one pectinate spiniform seta on distodorsal margin ( Fig. 11 N).

Pereopod-6 ( Fig. 11 F): Similar to pereopod-5, except basis with sensory-seta on sub-proximal dorsal margin. Propodus with three pectinate spiniform setae on distodorsal margin ( Fig. 11 O).

Pleopods ( Fig. 12 A): All five pairs similar, well-developed, biramous. Basal article, asetose. Rami lengths slightly more than twice width; proximal plumose seta on distal inner margin adjacent to articulation with basis. Endopod with inner and distal margins bearing 19−20 long plumose setae, distal most seta modified with whip-like tip; sub-distal lateral margin with seta modified with whip-like tip. Exopod with inner and distal margins bearing about 14−15 long plumose setae, outer margin naked.

Uropod ( Fig. 12 B): Biramous, about 1.1 times as long as wide, asetose. Exopod uniarticulate, with simple seta on mid-outer margin, and two simple setae (outer longest) on distal margin. Endopod biarticulated, article-1 slightly longer than exopod, with simple seta on sub-distal outer margin and two sensory-setae on distal inner margin; article-2 shorter than endopod article-1, with five simple setae of varying length on distal margin. Adult Male. Body ( Fig. 13 A): Length about 1.2 mm, about 5.2 times width.

Cephalothorax ( Fig. 13 A): About 20% of TL; sub-rectangular; with simple seta near each eye; longer than pereonites 1−3 combined. Eyes large, darkly pigmented with many ommatidia; diameter about half-length of carapace and about nine times of female eye.

Pereon ( Fig. 13 A): Slightly less 40% of TL, pereonites sub-rectangular, and wider than long; pereonites 1 to 6 progressively longer.

Pleon ( Fig. 13 A): Slightly over 30% of TL, as long as pereonites 2−6 combined; pleonites 1−4 subequal; pleonite-5 slightly longer than other pleonites.

Pleotelson ( Fig. 13 A): Little less than 10% of TL, length about two times width; slightly longer than length of pleonite-5, with eight (six lateral and two apical) simple setae on distal margin

Antennule ( Fig. 14 A): With seven articles, three peduncular and four flagella. Article-1 about 1.8 times as long as wide, with one sensory-seta on sub-distal dorsal margin, with three sensory-setae and one simple seta on ventral margin. Article-2 wider than long, about three times as long as article-1, with two sensory-setae and long simple seta on distoventral margin. Article-3 wider than long, with small seta on distodorsal margin and small simple seta on distolateral margin. Article-4 wider than long, with dense proximal group of aesthetascs ventrally. Article-5 about 2.1 times as long as wide, with distoventral row of aesthetascs (not illustrated). Article-6 about 2.3 times width, with distoventral row of aesthetasc (not illustrated). Article-7 about 3.7 times as long as wide, with one aesthetasc, one sensory-seta, and five simple setae of varying length, distally.

Antenna ( Fig. 14 B): Article-1 short, asetose. Article-2 about 1.1 times as wide as long, with long distodorsal simple seta; ventral margin forming shallow medial apophysis with weakly buttressed straight seta. Article-3 wider than long, with long distodorsal simple seta. Article-4, 2.3 times as long as wide, with two sensory-setae and two long simple setae of unequal length on distolateral margin; with two sensory-setae on mid-lateral ventral margin. Article-5, 2.7 times as long as wide, with long simple seta on distodorsal margin. Article-6 minute, with four simple setae of various lengths, two of them longer than articles 3 to 6 combined.

Mouthparts: reduced.

Maxilliped ( Fig. 14 C): Narrower than that of female. Basis with long simple seta near articulation of the palp, extending beyond palp article-2. Palp: article-1 asetose; article-2 triangular with small simple seta on outer proximal margin, inner margin with three (one long and two small) simple setae; article-3 with three simple setae of varying lengths; article-4 with five setae on distal margin, simple seta on outer sub-distal margin.

Cheliped ( Fig. 14 D −F): Coxal sclerite triangular, dorsally inserted, asetose. Basis about 2.3 times as long as wide, with simple seta on outer distodorsal margin. Merus triangular, with simple seta on mid-ventral margin. Carpus about 1.5 times as long as wide, dorsal margin with two small setae, one sub-proximal and one sub-distal; ventral mid-margin with two simple setae of varying lengths. Propodus about 1.2 times as long as wide, palm with inner face having “comb row” of ten stout setae just proximal to articulation with dactylus ( Fig. 14 F); fixed finger with crenulated ventral margin ( Fig. 14 D, F), with two ventral setae and three simple setae on outer incisive margin ( Fig. 14 D); stout slightly sinuate seta ( Fig. 14 E) on medial margin near articulation with dactylus ( Fig. 14 D). Dactylus with crenulated ventral margin, slightly longer than fixed finger, curved distally, with single dorsoproximal simple seta on inner margin ( Fig. 14 E).

Pereopod-1 ( Fig. 15 A): Longer than other pereopods. Coxa with simple seta. Basis slender, 4.4 times as long as wide, with dorso-proximal simple seta. Ischium wider than long, asetose. Merus, 2.7 times as long as wide, with two small ventrodistal spiniform setae. Carpus, 2.6 times as long as wide, with three short spiniform setae of unequal lengths on sub-distal margin. Propodus, 5.4 times as long as wide, with one sub-distal spiniform seta on ventral margin. Dactylus and unguis combined slightly longer than 2/3 of propodus, unguis longer than dactylus.

Pereopod-2 ( Fig. 15 B): Coxa with simple seta. Basis four times as long as wide, with dorso-proximal simple seta. Ischium wider than long, asetose. Merus about 1.7 times as long as wide, with ventrodistal small spiniform seta. Carpus about 1.9 times as long as wide, with one distodorsal and one distoventral small spiniform setae. Propodus about 4.6 times as long as wide, with one sub-distal spiniform seta on ventral margin. Dactylus and unguis combined slightly shorter than 2/3 of propodus, unguis longer than dactylus.

Pereopod-3 ( Fig. 15 C): Similar to pereopod-2, except basis to merus slightly shorter. Carpus slightly longer, with three distoventral small spiniform setae. Dactylus and unguis combined slightly longer than half-length of propodus.

Pereopod-4 ( Fig. 15 D): Basis slightly wider than in pereopod 1−3, with sensory-setae on mid-ventral margin, length about 2.1 width, with sensory-seta on sub-distal ventral margin. Ischium wider than long, with two ventral simple setae. Merus about 2.4 times as long as wide, with two spiniform setae distoventrally. Carpus about 2.3 times as long as wide, with simple distodorsal seta, and four stout bifurcate spiniform setae. Propodus about 4.1 times as long as wide, with distodorsal spiniform seta and with two spiniform setae on subdistal ventral margin, ventral margin with microtrichia. Dactylus and unguis combined shorter than propodus; dactylus longer than unguis.

Pereopod-5 ( Fig. 15 E): Similar to pereopod-4, except for basis and merus shorter. Propodus slightly longer.

Pereopod-6 ( Fig. 15 F): Similar to pereopod-4, except for basis longer. Propodus with three distal spiniform setae. Dactylus and unguis combined slightly longer than propodus.

Pleopod (not illustrated): Five similar pairs, similar to female but relatively more elongate with longer natatory setae.

Uropod ( Fig. 15 G): Biramous, similar to female, except for endopod article-1 having a row of transverse sensory-setae on inner mid-margin, and article-2 longer than endopod article-1. Narrower and longer in relation to pleotelson, terminating in long simple seta, slightly longer than pleopods 3 to 5 combined.

Remarks. The description for the male of A. hawaiensis represents the first for the genus Aparatanais . We did not observe any distinctive generic characters that would separate it from the limited number of males known for other related genera previously contained within Paratanais sensu lato.

For females the primary distinguishing generic character for all nine members of the Aparatanais is the presence of a stout dentate or spinulate spiniform seta on the inner margin of maxilliped palp article-2 ( Bird & Bamber 2013; Fig. 17 ). Preserved specimens of Aparatanais hawaiensis and three other species, A. denticulatus , A. lenoprimorum , and A. spinanotandus , exhibit a carapace divided into four apparent plates; a character not evident in the five remaining species to the genus. The presence of a relatively well-developed “kukri-shaped,” spiniform seta on the cheliped palm adjacent to the articulation with dactylus, distinguishes the new Hawaiian species from A. denticulatus and A. lenoprimorum , both of which apparently lack this distinctive character. The new species also differs from A. spinanotandus by having antennal article-2 with the dorsal margin bearing a dense row of minute setae.

The presence of a basally swollen, finely setulate seta on the epimeral margins of pleonites 1 to 4 distinguish A. hawaiensis , A. lenoprimorum , A. malignus , A. timutimu , A. tetradonta , and A. vicentetis , from A. denticulatus , A.

intermedius , and A. spinanotandus ( Bird & Bamber 2013) . The following key may be used to separate Aparatanais hawaiensis and the other five species characterized by having setulose setae on pleonites 1−4.

The taxonomic status of Aparatanais intermedius is uncertain, since it represents the only species within the genus that has the uropodal endopod fused into a single article and pleonites 1−4 without swollen lateral, setulose setae (see Dojiri & Sieg 1997), but until this information be corroborated by the examination of the type material its position within Aparatanais remains questionable ( Bird & Bamber 2013).

Figures

FIGURE 8. Aparatanais hawaiensis sp. nov. holotype female. A, dorsal view; B, lateral view; C, enlargement of basally swollen setulose seta on pleonites 1 – 4. Scale bar = 1.0 mm for A – B.

FIGURE 9. Aparatanais hawaiensis sp. nov. holotype female. A, antennule; B, antenna; C, labrum; D, left mandible; E, lacinia mobilis; F, right mandible; G, labium; H, maxillule; I, maxilla. Scale bars = 0.1 mm for A – D and F – I.

FIGURE 10. Aparatanais hawaiensis sp. nov. holotype female. A, maxilliped; B, enlargement of palp article- 2; C, right cheliped, lateral view; D, enlargement of kukri-shaped sinuate spiniform seta; E, right cheliped, inner view; F, enlargement of setulate setae. Scale bars = 0.1 mm for A, C and E.

FIGURE 11. Aparatanais hawaiensis sp. nov. holotype female. A − F, pereopods 1 – 6; G – O, enlargement of different kind of setae and spiniform setae in pereopods. Scale bars = 0.1 mm for A – F.

FIGURE 12. Aparatanais hawaiensis sp. nov. holotype female. A, pleopod; B, uropod. Scale bars = 0.1 mm.

FIGURE 13. Aparatanais hawaiensis sp. nov. paratype adult male. Dorsal view. Scale bar = 1.0 mm.

FIGURE 14. Aparatanais hawaiensis sp. nov. paratype adult male. A, antennule, B, antenna, C, maxilliped; D, cheliped, lateral view; E, enlargement of stout slightly sinuate seta; F, propodus and dactylus, inner view. Scale bars = 0.1 mm for A – C and D.

FIGURE 15. Aparatanais hawaiensis sp. nov. paratype adult male. A − F, pereopods 1 – 6. G, uropod. Scale bar = 0.1 mm.

FIGURE 16. Digital images of Aparatanais hawaiensis sp. nov. holotype female: A, dorsal view of habitus; B, lateral view of habitus. Scale bar = 1.0 mm.

FIGURE 17. Palp article- 2 of maxilliped + enlargement of the spiniform seta: A, Aparatanais denticulatus; B, A. hawaiensis sp. nov.; C, A. intermedius; D, A. lenoprimorum; E, A. malignus; F, A. spinanotandus; G, A. tetradonta; H, A. timutimu; I, A. vicentetis. Figures modified from: A, Guţu & Ramos (1995); B, Morales-Núñez, Pelleteri & Heard (this study); C, Dojiri & Sieg (1997); D, Tzeng & Hsueh (2014); E, H, Bird & Bamber (2013); F, Sieg (1981); G, Bird (2015); I, Larsen (2012). Not to scale.