Dicamptodon copei, n. sp., from The Pacific Northwest, U.S.A. (Amphibia: Caudata: Ambystomatidae) Rowaup A, Nusspaun Dicamptodon copei described from western Washington State and northwestern Oregon. It is characterized by sexual maturity in the larval sage at small size, short limbs, small head, reduced tail fin, reduced number of maxillary + premaxillary cecth, reduced number of vomerine teeth, and reduced sens y to thyroid treatment; as compared to larvae of Dicamptodon ensatus, It is suggested that D. coped is a relatively recent derivative of a D. ensatuclike ancestor. TUDIES of geographic variation in Di camptodon ensatus have shown that some populations are dimorphic tion of the metamorphic processes and re- productive biology af each morph has led ime to conclude that twe species are involved. One is the well established D. ensatus (Eschschaltz); the other is an unnamed species described below, Examina- Dicamptodon copei h. sp. Figs. 1.2 Holotype.—U. §, National Museum 166784 (Fig. 2), breeding, branchiate male 92.2 mm snoutvent length (live measurement from snout to anterior corner of vent}, collected by Ronald A. Nussbaum 20 June 1969 in Mar(rjatta Creek 85 m upstream from bridge on state highway 504, SW Sec. $, T.9 N., R. 4 E,, 45° 17° N, 122° 18” W, 810 m eleva- tion, Cowlitz County, Washington, Paratypes—USNM 166785-1668]4, 15. ma- ture branchiate females and 15 mature branchiate males 68,9-105,8 mm (x 84.9 mm) snout-vent length, collected by Ronald A. Nussbaum at the typelacality, Diegnosis—A small, attenuate species of Dicamptodon known only from the larval stage, Differs from D. ensatus by having a narrower head (head width 018-021 (3 0.194) times snounvent Iength as opposed to O.21-0.28 (2 0.228) in ensntus); by having, a shorter head (head length 0.39-0.49 (¢ 0.431) times body length as opposed to 0.46- 0.55 (é 0.504) in ensatus}; by having shorver limbs (coupling value 0.83-1.10 (& 0.968) for copei and 0.79-0.98 (€ 0.863) for ensatus): by having more costal folds between ad- pressed limbs (00-20 for copei and 0.5 10 overlapping by 1.0 in ensaits); by having lower tail fins (maximum tail height 0.18-0.28 {% 0.196) times snout-vent length as compared to 021-027 (& 0.238) for ensatus); by having shorter tail fins (usually extend anteriorly on the back to a point which is posterior 40 the vent in copet and opposite to or an. terior to the vent in ensains); by having fewer maxillary 4+ premaxillary teeth (88-44 {& 89.5) as compared to 46-68 (8 584) in eusatus); by having fewer vomerine teeth (28-42 (7 84.4) as compared to 41-59 (F 47.5) in enwatus); by having no palatopterygoid teeth (palatopterygoid teeth may or may not be present in ensalus larvae); by having fewer gill rakers on the anteriormost two rows and the posteriormost two rows (see Tables 1, 2). In coloration, D. copei has less dark mottling on the tail fin at all sizes than does B, eusatus. The venter of cape’ is generally much darker than sympatric ensutus at sizes greater than 50 mm. snout vent length, Conspicuous granular glands are seen on the dorsum of copei (Fig. 2) these are less apparent in ensatus. D. copet is characterized by sexual maturity at small sizes (mature paratypes 689-1038 mm (x 84.5 mm) snautvent length) as compared. to sympatric, branchiate ensatus which ma- ture at sizes greater then [20 mm snout vent length. D. copei is less responsive to thytoxine than is ersaéus. Dewription of halotype—Male, left testis white, bilobed, anterior lobe 115 mm, posterior lobe 9x4 mm; cloacal glands swollen. Proportions: (all measurements taken afver 48 hr fixation in 10% formalin) snout to anterior edge of vent 88.4 mm; snout to posterior edge of vent 92.5 mm: anterior edge of vent to tip of tail 718 mm; head length 26.4 mm; head wideh 164 mm; right forelimb 22.5 mm; left forelimb 506NUSSBAUM—NEW DICAMPTODON FROM THE PACIFIC NORTHWEST, 507 Fig. 1. Dicarptodon cope, showing extent of metamorphic changes after months of thyroid twcativent. Adult color fs tagely contined to Dead. (LIS¥). me 282 mm; right hindlimb 26.8 mm; left hindlimb 26.7 mm; digits in’ order from Jongest to shortest, on forelimbs $2.14, on hindlimbs $425-1, axilla to groin distance 450. mm; maximum tail height 15.9 mm: distance beween medial edges of openings to external nares 6.7 mm: distance between anterior corners of orbits 7.3 mm; distance from anterior corner of right orbit to right external naris 89 mm; distance from an- tetior comer of left orbit to left external naris 4.3 mm: length of right orbit 3.0 mm: length of left orbit 8.0 mm, Counts: num ber of costal grooves right and left 13; Fig. 2 The typespecimen of Dicamplodon copei (upper) compared to a larval D. ensetus (lower) of similar mout-vent length also collected at the type-locality of D. coped. Note more slender propor- tions, smaller head, and. promi sat skin glands in D. copel. (0.8%),508 COPEIA, 1970, NO. § Tamie 1, Comramsox op Grr. Raker Counts Berwesw Dicamplodon ensatus ano D. capei. D. ensatus (n= 20) 355% Vicoeral Na. Gath Relation Confidence Arch Bakes Frequency ‘Taterval ard 7 O2t 012-046 (posterior edge} 6 ass 0.40.88 4th T a0 0.02-0.97 fanterior edge} 5 083 0.65-0.98 5 aor o.01-0.22 ath 4 007 010.22 035 015-059 (posterior edge) 8 80 61-092 0.00 0.35-0.81 5 043 004-031 1.05 000-025 Sth 7 000 0.00-0.12 025 008-049 (anterior edge) 6 0.80 61-092 a7 051-041 5 080 008-039 000 000-017 Sth 6 007 oo1-n2e 05 051-091 (posterior edge) 5 090 073-098 0.20 0,060.44 4 003 0-017 0.05 0.00-0.25 oth 5 om 0.00-0.17 070 a6-0.88 (anterior edge) 4 og 0.78-0.90 030 12-054 3 os 000-0.17 0.00 0.00-0.17 costal folds between adpressed limbs right and left sides 6.5; number of maxillary + premaxillary teeth 39; number of vomerine tecth, right series 19, left series 22; number of dentary teeth 51; six rows of gill rakers on either side, number of gill rakers per row anteriormost to posteriormast row, right side 6.65.65-4, left side 66-66-55. Color in life: dorsum brown with patches of yellowish tan overlying conspicuous granular glands, scat- tered coppery iridophores barely discernible to unaided eye; sides grayish brown with patches of yellowish tan pigment as on dorsum, seattered coppery iridophores; venter dark bluish gray, effect caused by numerous small white openings to skin glands sur. rounded by melanophores; tail brown along the sides with yellowish tan patches, es pecially overlying glandular areas at base of dorsal fin, gray below with scattered patches of yellow, lips of cloaca white; limbs brown above with patches of yellowish tan, colored like venter below, black callosi tips of digits; head brown above with yel- lowish tan patches and scattered coppery itidophores, concentration af cuppery irido- phorcs decp-lying above each cye, throat and chin dark bluish gray like venter bur grading tw brown on margins of jaw: eye, pupil surrounded by narrow ring of yellow- gold pigment invaded with coppery irido- phores, faint seattered irregular patches of yellow-green painted an surface of iris, scat- tered flecks of yellow.gold pigment. above and below in iris separated by irregular horizontal brown band; gills dark purple, fimbriac with scattered melanophores and yellow lipophores. Range.—Forms referable wo D, copei have been collected at the following local Wastunorox: (ributaries of the Quinault River, Jefferson and Grays Harbor counties; tributaries of the Hoh River, Jefterson Go, near Lake Cushman, Mason Co; Sol Duc Falls, Clallam Co,: near a fish hatchery east of Vancouver, Clark Co.; Elk Creek, Cowlitz Go. tributaries of the White Salmon River, Skamania Co.: Onzcon: in two smail tribu- taries of the Galumbia River, Multnomah Co, In general, it appears that D. cope’ is restricted to western Washington induding the Olympic Peninsula and. the southern Washington Cascade Mountains. Apparently it barely enters Oregon in the Columbia River Gorge. The distributions of D. coped and D. ensatus are presently being studied, andNUSSBAUM-NEW DIGAMPTODON FROM THE PACIFIC NORTHWEST 509 Tante 2. COMPARGON OF BRANCHIATE Dicamptaden ensatus To 1, copei; Mean + Two STANDARD ‘Eanoes, by Store, a= ¥ Inveacert, r = Conretarion Corrricren. D, ensatue (x = 80) D. copei (an = 30) vomerine teeth 405 CE 2155) HA ( L500) range = 41-39 range = 28-42 maxillary + pre- 58.4 (= LM) 995 ( 1.088) maxillary teeth range = 46-68 range = 33-4 coupling value 0.863 (O18) 0.268 (= 0.020) range = 0.789-0.994 range = 0.891=1,064 head length b = 0.885 (+ 0.088) hh = 0.275 (= 0054) regressed! on a = 6412 (= 2.150) a GIBL (% 3252) body length r= 0879 r= 0.888 bhead. width b = 0.290 ( 0.022) b = 0219 ( 0.028) reweessedl on a = 2493 (= 124) a = 3506 ( 2551) body length F = 0987 F = 0.902 maximum tail b = O48 (x anes) b = 0.061 ( 0.058) height regressed on a = 4088 (= 2.468) a = 11574 (+ 5.070) anoutvent length r= 0950 1 = 0306 hindlimb length 0485 CE 0.050) b = 0.299 (= O.o48y sregresied on 1.58% Ge 2811) a = 7.598 (& BASH body length = 0972 r= 0918 forelimb | 0355 (= 0.088) ‘b= 0.279 (= O04) regressed on) 2588 ( 2.160) a = ATS (® 2640) body lengtis 0.975 r= 0982 the Tanges of the two species will be dis. cussed in more detail in a forthcoming paper. Etymology.—The specific name copei is a patronym honoring Edward Drinker Cope for his many contributions to our knowledge and understanding of vertebrate morphology and phylogeny. Suggested common name—I recommend this new species be called Cope’s salamander. Companison of D. copei to D. ensatus Dicamptodon copei occurs with D. ensatut at the type-locality, although the former is present in greatest abundance, This sym- patry made recognition of two species pos sible early during the course of investigation. The comparisons to follow are based on the 30 paratypes of D, copei and 20 larval D. ensatus of similar size, from the same Site and sexual maturity.—D. ensatus larvae may metamorphose into terrestrial adults during or following their second year, or become breeding Tarvae at large size. If metamorphosis occurs usually at sizes from 54 t 93 mm snoutvent length (Nuss- baum and Clothier, in prep.). Size at sexual maturity varies geographically in D, ensatus larvae, although it usually occurs at sizes greater than 115 mm snoutvent length. At the typelocalicy of D. copei, larvae of D. ensatus are not sexually maturé at sizes up to 120 mm snoutvent length, but at these sizes they are well on their way to heeoming breeding larvae and are not likely to meta morphose. D. copei larvae attain sexual maturity at small size. The 30 paratypes, ranging from, 69 to 104 mm snoutvent length, are all sexually mature, The males have wellde- veloped testes, and the cloacal glands are swollen in most. Some females have well- developed ova ranging from 2.1 to 5.5 mm in diameter and swollen oviduets. A few of the female paratypes had oviposited previous to capture as evidence by the greatly swollen, twisted mature of the oviducts and flaccid, loose appearance of the ovaries which con tain small, yolky developing ova, ‘Metamorphosis Metamorphosis was in- duced in order to determine the color and pattern of adults which ate not always510. HEAD LENGTH (mm) 40 50 60 70 BODY LENGTH (mm) Fig. 5. Head length regressed on body length. Open circles=Dicamptodon ensatus, closed circles =D. copei. Lines fitted by least squares method. See Table 2. 80 available by colledting, It was noted in 1967 that some Maratta Creek larvae did not develop a full color pattern when treated with powdered thyroid, A control experi- ment was begun 22 August 1968 to deter- mine the metamorphic relationships of the two forms from Maratta Crock. Two aquaria, exch containing both types of Dicamptoden were placed in a controlled temperature room, Small amounts of powdeted thyroid were added to the water in both aquaria ‘The thermal histories of all the larvae were the same, their sizes were comparable, and because each container keld both types of larvae, the dosage and length of exposure to thyroxine were the sime for both types of larvae within each aquarium, The only difference inherent in the experiment was that D. coped were sexually mature and D. ensatus immature over the sire range used. However, immature D. coped teated sepa. rately, reacted much the same to thyroid treatment as did mature D, copei. Experi- ments have shown {date to be published elsewhere} that large sexually mature D. ensatus larvae are slightly less reactive to thyroxine than smaller immature D. ensatus, But even extremely large D. ensatus lar (170-180 mm SV) are much more sensitive than mature D. coprei ip holds for D. ensatus from the typelocality of D. capei as well as from other localiti Metamorphosis was rapid in 2D. ensafus in both aquaria, and comparable to metamor- 1970, NO. HEAD WIDTH (mm) 40 50 60 7 80 BODY LENGTH (mm) Fig. 4. Head width regressed om body length Open creles=Dieumpodon ensatus, closed citcles =D. cope. Lines fitted by least squares method. See Table 2. phosis of D. ensatus larvae from other pares of the range. D. ensatus had full color and pattern after three months at 10° C, Several D. ensatus larvac from Maratta Greek held in separate containers initiated and com- pleted metamorphosis without thyroid treat- mént. It is evident that a changed environ- ment can induce metamorphosis in species. Metamorphie changes were much slower in the treated D. capef. Reduction of fi gills, and lip folds occurred normally but at a slower rate; eye protrusion and color de- velopment were delayed even more, Five months elapsed before any adult color ap- peared, and this was confined to the snout region. At the end of 11 months the adule color pattern had spread from the tip of the snout posteriorly on the head, and spots of color had appeared on the upper eyelids and on other parts of the body, bur only to a slight extent (Fig. 1). ‘The develop- ing pattern af color in D. coped is similar to that of D. ensatus but apparently does not go to completion. Alter 11 months, treated D. copei had developed well-protruded eyes and fully formed eyelids. Color compariens—At 50 mm snout-vent, both species have white venters with melano- phores encroaching from the sides, At 90 mm, melanophores have completely en- circled the body in both species, but melano- phores are generally denser on the venter of D. cope: giving it * darker appearance.NUSSBAUM—NEW DICAMPTODON FROM THE PACIFIC NORTHWEST ou Fig. 5. Tails of Dicamptodon ensatus (upper) and D. copei (lower). Tail fin of D. copel re duced in sie. (078x). Young of both species have light mottling on the upper surfaces of the tail fin. The ‘mottling, is indistinet in D. cope’ and promi- nent in D. ensatus. In addition, young D. encatus larvae have a blacktipped tail young D. copei lack the black tip. Tai mottling fades in older individuals of both species, as does the black tail tip in D. ensatus. The dorsum and sides of young and old D. copei_have prominent yellowish an patches (Fig. 2) covering aggregations of large, white granular glands which are casily visible to. the unaided eye. Faint patches of yellowish tan can be seen in young D. ensatus larvae, but are more difficult to discern in older larvae. D. ensatus larvae of all sies lack the enlarged granular glands seen in D. copei. Some populations of D. ‘encatus, especially in southern Oregon and northern California, have light markings on the dorsum and sides which superficially resemble those found in D. copei. However, the enlarged granular glands found in D. copet are not present beneath these light markings. Also, the markings found in non- sympatric D. ensatus vary from extremely large blotches to fine streaks, totally unlike the pattern seen in D. cope. Body proportions—The 30 paratypes of D. copei were compared w 20 larval D. ensatus of similar size collected at the type- locality of D. copei. ‘The following measure- ‘ments were taken from each specimen: snout-vent length, from tip of snout to an- terior edge of vents head length, from tip of snout to 2 point just above the insertion of the gills where the jaw muscles form an angle with the neck (this measurement has proven (o have a smaller coefficient of varia- tion than the midline distance from the tip fof the snout to the cage of the gular fold) head width, taken at the angle of the jaws; axillagroin disance; forelimb length, from tip of longest digit to body wall hindlimb length, from tip of longest toe to body wall: ‘maximum tail height; tail length was also taken but not used because some specimens had regenerated tail tips. Body length i snoutvent length minus head length. ead length and head width regressed on ody lenge (Figs 8, 4; Table 2) show that D. copei have significantly shorter and nar- rower heads than D. ensatus. ‘This difference is apparent by simply looking at the animals (ig. 2) Regression of forelimb and hindlimb on body Tength show that both limbs are sig: nificantly shorter in D. copel (Table 2) “The coupling values (exila-groin forelimb -+ lim) for the ewo species are significantly MAXIMUM TAIL HEIGHT (>) ht SNOUT-VENT LENGTH (rm) Fig. 6 Maximum tail height regressed on snoutvent “Open. circlet = Dieampiodon ensatus, closed circles =D. copel. Lines fitted Dy least squares method. See Tabie 2.sig ig. 7. Comparison of transformed Di Both collected as larvae at type-locality ot COPEIA, 1970, NO. 8 intadon ews (pen a astomed Doped Comer ‘small head and lack of color in D. copei. Animals have been anesthetized and left ential thyroid treatments. Note be oF Both ‘arranged to demonstrate the shorter legs of D. cope. (0.78%). different (Table 2), reflecting the shorter limbs in D. copei. ‘The tail fin of D. ensatus is visibly higher than the tail fin of D. cope’ (Figs. 5, 6; Table 2), and usually extends farther an teriorly on the back, ‘The reduced tail fin is noticeable even in small, immature D. copel Counts—The number of costal grooves was the same for all specimens of both species. By counting one cach in the axilla and groin the number was 18. However, the posteriormost groove starts above the hind- limb and curves forward into the groin; some investigators would not count this groove and would report 12 costal grooves. By my method of counting, 18 costal folds corresponds with 14 trunk vertebrae as shown through the use of radiographs. ‘The number of costal folds between ad- pressed limbs, estimated to the nearest half, was taken for all specimens. ‘This character reflects limb length in this case (as both species have 14 trunk vertebrae), but is included here because it is a better key character than absolute limb length or the coupling value. D. copei has 0.0-20 costal folds between its adpressed limbs, while the limbs of D. ensaius usually overlap or meet (Figs. 7, 8). ‘The last four visceral arches of branchiate Dicampiodon have rows of gill rakers. The third arch has one row of gill rakers on the posterior edge. ‘The fourth and fifth arches have two rows, one on their anterior and ‘one on their posterior edges. The sixth arch hhas a single row of gill rakers on the anterior ‘dge. Gill raker counts were made for all six rows. Table I shows that D. ensatus has 2 higher number of gill rakers than D. cope’ ‘on the two anteriormost and the two pos teriormost rows. Premaxillary and maxillary veeth were counted as a single unit. The number of maxillary plus premaxillary teeth is sig- nificantly higher for D. ensatus (Fig. 9: Table 2) D. ensatus also haye more vomerine teeth than D. copei (Fig. 10; Table 2). In addition, the vomerine series in D. ensatus extends t0 ‘or beyond the posterior margins of the COSTAL FOLDS BETWEEN ADPRESSED LIMBS. 8 ‘SNOUT VENT LENGTH (7) Fig. 8. Costal folds between adpressed limbs. Oped cnces Skies =D: cope dlosed jcarmptodom ensatus,NUSSBAUM—NEW DICAMPTODON FROM ‘THI MAXILLARY + PREMAXILLARY TEETH % bo io zo SNOUT -VENT LENGTH (mm) ig. 9. Manllary + premanillary teth in larva of Dicamptadon ensatus (open circles} and D. copei (closed circles). See ‘Table 2. choanae, while in D. copei the vomerine series extends posteriorly only to the middle of the choanae, or slightly past the mi point. Six of the 20D. ensatus hawe a small series af teeth posterior to the vomerine serics associated with the palatopterygoid on both sides, eight have this series on one side only, and six of the 20 lack this series en- tirely. There are $8 tecth on each af the palatopterygoid series. They were not counted with the vomerine teeth although they are in line with them. None of the $0 paratypes of D. cope’ has a palatopterygoid series on either side, Discussion Presently it is not known whether D, copei ever metamorphose in nature. No adults have been found at the typelocality despite many hours of search, D, ensatus may wrans- form at the typelocality of D. coped as several wransforming larvae and one fully transformed adult have been collected there (the latter collected by Dr. Dean E, Metter, University of Missou Sexual maturity at small size and delayed and incomplete reaction to thyroxine sug- gest that D. copei may nat metamorphose in nature. This prospect presents problems which are difficult to reconcile. Obligate neoteny in salamanders limits the means of dispersal and therefore creates problems of survival in rapidly changing environments. Most permanently aquatic species of sala- manders oceur in geologically olel areas such as Appalachia in eastern North America. PACIFIC NORTHWEST 513. VOMERINE TEETH SNOUT = ENT LENGTH (em) Fig. 18, Vomerine teeth of larval Dicomp- torton ensatus (open circles} and D. copeé (closed circles). ‘See Table 2 In suels regions, drainage systems and under- ground waterways have stabilized, whieh in some respects eliminates the necessity of dispersal at least until the environment be- comes unstable again, The relatively recent mountain systems of western North American do not offer stability, and major river systems and drainage patterns are still being formed. ‘The establishment of an obligate neotene under such conditions seems unlikely, The facultatively neotenic D. ensatus would seem to be better adapted wy such environments, It would, therefore, not be surprising if further investigation shows that D. copei do metamorphose under certain environ- mental conditions or at different localities, It is likely that D. copet has evolved from an ensatuslike ancestor, and during rela- tively recent times. The differences berween larval D. ensatus and D. copei are not great. Fo derive D. copei trom D. ensalus requires only that the reproductive organs mature at small body size; that differential allometric growth patterns produce smaller heads (and this may account for the reduced number of teeth), shorter limbs and reduced tail fins, and that enlarged acinous glands develop at small body size, These and other changes easily could have evolved through paedomor- phosis and heterchrony, processes which are known to have played prominent roles in salamander evolution. This type of evolu: tion apparently can occur rapidly and does not require extensive genetic reconstruction. No hybrids between D. cope? and D. ensarus have been found, Isolating mech- anisms are presently unknown, althoughSlt size may be a factor. Large D. ensacus larvae are highly cannibalistic. At the typeltocalicy of D. copei, stomachs of large neotenic D. ensatus commonly comiain immature D, ensatus larvae as well as sexually mature D. coped. It is suggested that size may in part determine whether an individual is a potential mate or food ites ‘The typelocality af D. cope’ is situated ‘on the western slopes of the Cascade Range, an area which receives excesive precipita: tion from moistureladen air masses moving eastward from the Pacific Ocean. south and eastward precipitation is lower, but is high enough to meet the requirements of the more wide ranging D. ensatus. Coastal Washington has probably been an area of rainfall since its origin. During the Pleistocene pluvials, precipitation in coastal Washington was much higher than itis taday; and much of the land was below the snow and ice zone which capped the Cascade Range during the periodic ice advances. Dur- ing glacial maxima, coastal Washington must have acted as a refugium for mount dwelling species. Dumas (196) has suggested that Rana cascadae evolved from ancestral Rana pretiosa wi laced in this hypo- theti¢al refugium in Pleistocene time. Such a theory would also best explain the relation- ship between the two species of Dicamptodon, Perhaps a segment of ancestral D. ensatus became isolated in coastal Washitgton dur- ing an early glacial maximum. Evolution toward obligate neoteny could have pro- ceeded rapidly because the advantages of terrestrial dispersal were reduced in such an ‘To the COPEIA, 1970, NO. $ area with excessive rainfall and permanent lakes and streams. The sympatry of D. ensatus and D. capei at the type-locality and in the Columbia River Gorge may be a result of adjustments in the ranges of both species during postglacial time. ‘This theory requires 2 more rapid rate of speciation than is usually invoked. However, considering the type af evolution the close relationship of the two species, and dhe strong selective forces asiociated wi extremes of climate, the proposal gains credence, A more complete study of the relationships bevween DP, ensatus and D. copei, and an analysis of geographic variation between populations of Dicamptadon from through: ‘out the range, is in progress. ACKNOWLEDGMENTS. T thank Dr, Robert M. Storm for reviewing the manuscript, Dr. Edmund D. Brodie Jr. for photographing the specimens (Figs. 1, 2, 7) and for reading the manuscript, Dr. Philip C. Dumas and Dr, Robert H. Brown for encouragement and advice early during the course of my studies of variation in camptodon, and Dr. Dean E, Metter for the benefit of his field experience. Liveratur Grrep Duss, P. C. 1966. Studies of the Rana com plex in the Pacific Norehwest. Copeia 1906(1) 74. DEPARTMENT OF ZooLocY, ORECON STATE Univexsrry, Corvarus, Orecon 97331.