Pinnotheres novaezelandiae Filhol

Page, Roderic D. M., 2012, Description of a new species of Pinnotheres, and redescription of P. novaezelandiae (Brachyura: Pinnotheridae), New Zealand Journal of Zoology 10 (2), pp. 151-162 : 152-158

Pinnotheres novaezelandiae Filhol

( Fig. 1, 2 A-H)

Pinnotheres pisum (Linnaeus, 1767) . Heller, 1868: 67. ­ Miers, 1876: 48. - Filhol, 1885a: 50; 1885b: 394. ­ Adensamer, 1897: 105-106 (part). -Thomson in Ayson, 1900: 22. - Hutton, 1904: 250. -? Chilton, 1911: 255-256. - Thomson, 1913: 238. - Borradaile, 1916: 100-101 (part; not text-fig. 12 fide Bennett (1964, p. 76». - Tesch, 1918: 249, 251, 287. ­ Thomson & Anderton, 1921: 101. -? Gurney, 1924: 195 (part). - Chilton & Bennett, 1929: 775-776. ­ Richardson, 1949: 30, 36, fig. 23.

Pinnotheres novaezelandiae Filhol, 1885a: 15-16 , 50; 1885b: 395-396, 495, pl. 46 fig. 1--ti. -" Lenz, 1901: 467, pl. 32 fig. 11-14. -" Hutton, 1904: 250. ­ " Tesch, 1918: 249,251,287.-""oung, 1929: 152.­ Powell, 1937: 371, 382; 1947: 39-40. -" Richardson, 1949: 36. - Scott, 1961: 303-309, fig. 1, 4, and 6 (part). - Dell, 1963: 50, 2 figs. - Bennett, 1964: 76­ 79, fig. 84,85,88,92, and 93 (part). -" Wear, 1965: 16, 18, fig. 6D. -"Silas & Algarswami, 1967: 1175, 1203,1207. -" Morgans, 1967: 171. - Dell, 1968a: 26 (part); 1968b: 233, 238. - Morton & Miller, 1968: 292 (part; not fig. 101). - Schmitt et al., 1973: 58-59 (part). - Hayward, 1974: 159, fig. 1 and 2. ­ Carpenter, 1976: 16. -? Gordon & Ballantine, 1976: 122. - Jones, 1977: 145-158,fig. 1-7; 1978: 667--ti76. - Fenwick, 1978: 208. - Hickman, 1978: 211-215. ­ Pregenzer, 1979: 22-30, fig. 4, 9, 12, 20, 20A, 28, and 29. - Jenkins, 1979: 44-45. - Baxter, 1982: 77­ 83.

Pinnotheres schauinslandi Lenz, 1901: 468-469 , pl. 32 fig. 15-18. - Chilton, 1911:295-296. - Tesch, 1918: 250­ 251, 287. - Chilton & Bennett, 1929: 775-776. ­ Scott, 1961: 303, 307. - Bennett, 1964: 79 --80, fig. 87 and 89-91 (part; not fig. 86). -Silas & Algarswami, 1967: 1209, 1221. - Schmitt et al., 1973: 85 (part).

? Pinnotheres sp , Dell, 1960: 5.

Pinnotheres sp. Jenkins, 1979: 44-45 , fig. 46.

Not Pinnotheres novaezelandiae Filho 1. -" Chilton, 1911: 295-296. - Takeda & Miyake, 1969: 180-181. (= Pinnotheres atrinicola n.sp.)

Not Pinnotheres novaezelandiae Filhol. - Rathbun, 1923: 98, text-fig. 2, pl. 16 fig. 2. -" Guiler, 1952: 40. (= Pinnotheres hickmani (Guiler)

Diagnosis. Chelae of hard-stage without a continuous dorsal row of setae on propodus. Terminal segment of hard-stage abdomen trapeziform. Male 1st pleopod stout, blade-like, its distal end slightly curved, densely setose. Legs of mature female not markedly asymmetrical. Second leg with dactylus always shorter than carpus, carpus and propodus subequal in length. First 3 legs without long setae on carpus, propodus, and dactylus.

First-stage zoea without dorsal and lateral carapace spines; rostrum minute. Lateral carapacial chromatophore with a single centre; black subin-

testinal chromatophore in telson proximal to yellow lateral intestinal chromatophore.

Description. HARD-STAGE (Fig. lA-G). Carapace ( Fig. 1A) hard, smooth except for minutely pitted frontal and branchial regions, oval to circular, strongly arched longitudinally, convex dorsally; width range 3.4-11.3 mm; lateral margins nearly vertical; front protruding, deflexed downwards, convex or straight in dorsal view, sometimes with a medial notch. Eyes visible in dorsal view.

Third maxilliped endopod ( Fig. 1B) with ischium and merus fused; aboral surface convex, with small setae scattered over surface; lateral margin with a dense row of stout setae; medial margin and medial oral surface with long, plumose setae; medial aboral surface with a row of stout setae; lateral margin of carpus and propodus with a dense row of cu~ed setae; dactylus arising near propodus/carpus articulation; distal end of dactylus bearing long setae. Chelipeds stout; merus with a row of setae on dorsal and ventral margins; carpus with a row of setae on ventral margin and a dense tuft of setae dorsally on inner face; chelae ( Fig. 1 C,D) stout; propodus inflated; inner ventral margin ( Fig. 1D) with a row of distally directed setae extending from articulation with carpus to tip of fixed finger; propodus with tufts of setae dorsally and ventrally near articulation with carpus, a short row of setae on outer face below articulation with moving finger ( Fig. 1C), and a short row of setae on inner dorsal face at articulation with moving finger ( Fig. 1D), extending on to proximal half of moving finger; fingers stout, crossing when closed, each with a proximal tooth; teeth biting together when fingers closed; a single row of stout, dagger-like setae between tooth and tip on both fingers, flanked on either side by longer setae; a short row of daggerlike setae on inner face of fixed finger immediately proximal to tip ( Fig. 1D).

Legs flattened, the 4th shortest, the 2nd and 3rd subequal in length. Merus of each leg bearing a row of long setae on dorsal margin; a dense fringe of setae on ventral margin of each merus, carpus, and propodus; carpus of each leg with a tuft of short setae dorsally at articulation with propodus; 2nd and 3rd legs each with 2 rows of long, pinnate setae, one on ventral margin and one on posterior face near dorsal margin of last 3 segments; merus and dorsal and ventral margins of carpus and propodus of last leg with similar setae (these long setae reduced or absent in large males).

Abdomen 7-segmented. Male abdomen (Fig. IE) tapering from segment 3, the broadest, to segment 7; margins of 3rd segment markedly co?vex; terminal segment trapeziform; segments fnnged with short setae. Female abdomen (Fig. IF) broader and less tapering than in male; 2nd segment

relatively broader; terminal segment more rounded.

Male 1st pleopod ( Fig. 1 G) densely setose, stout, blade-like, tapering gradually, its distal end slightly curved laterally; medial row of setae terminating on sternal side of pleopod just below tip.

Colour pattern complex, clearly defined, strongly pigmented. Anterior half of carapace orangebrown, with white spots and cream areas forming a distinctive pattern ( Fig. 1A); posterior half more variable in pattern, with white spots and markings on a mauve and yellow background. Legs brown with darker areas; chelae yellowish-brown with brown markings. Thoracic sternites creamy yellow with brown patches along junctions between individual sternites. Abdomen with paired brown patches dorsally. Both the colours and their extent are variable.

MATURE FEMALE ( Fig. 1 H-J). Carapace (Fig. IH) soft, smooth, longitudinally arched, dorsally convex, oval in dorsal view, laterally angular; width range 7.8-20.0 mm; front rounded. Eyes not usually visible in dorsal view.

Third maxilliped (Fig. 11) as for hard-stage, but lateral setae on fused merus/ischium and on carpus shorter and sparser.

Cheliped merus with a short row of setae dorsally ~nd another ventrally, the latter row reduced or absent in large females; carpus with a row of setae dorsally on inner face; chelae stout, variable in shape, the propodus glabrous except for a row of setae on ventral face; setation of biting surfaces of fingers as for hard-stage; inner tip of moving finger with a few setae.

Legs subcylindrical, the 4th shortest, the 2nd and 3rd subequal. Second leg (Fig. 11) with carpus and propodus subequal (length ratio of carpus to propodus 1:1.09 j: 0.07 SD; n = 28), dactylus always shorter than carpus. Legs on each side symmetrical (ratio of sum of lengths of last 3 segments of 2nd leg, shortest to longest, 1:1.02 j: 0.02 SD; n = 27). Merus of each leg with a row of long setae along dorsal margin. First 3 legs with scattered setae on ventral margin of propodus and dactylus. All 4 legs with dactylus bearing a row of fine, tooth-like setae on ventral margin.

Abdomen 7-segmented, as wide as carapace or wider, its edges usually extending to coxae of legs; segments fringed with setae.

Colour pattern (where present) as for hard-stage, but less heavily pigmented, and pattern less well defined; otherwise creamy white all over. Mature ovaries clearly visible through carapace as a purple mass.

FIRST-STAGE ZOEA ( Fig. 2 A-H). Carapace ( Fig. 2A) globulose, without dorsal or lateral spines but with a pair of setae dorsally; mean length 0.50 mm (range 0.45-0.55 rnm), mean width 0.37 mrn (range 0.35­ 0.38 mrn). Rostrum minute, variable in size. Antennule ( Fig. 2A,B) a small, hemispherical bud with 2 aesthetascs and 1 seta. Antenna ( Fig. 2A) minute, bearing 1 seta. Mandible: incisor process ( Fig. 2C) stout, with a major tooth and 2 smaller accessory teeth.

Maxillule ( Fig. 2D): endopod of 2 segments, with 4 (0/4) setae; basal endite with 5 setae; coxal endite with 4 setae. Maxilla ( Fig. 2 £): endopod with 3 setae; bilobed basal endite with 4 and 5 setae; coxal endite with 5 or 6 setae, the 6th (where present) minute; scaphognathite with 4 plumose setae and a plumose proximal extension.

Maxilliped 1 ( Fig. 2F): basis with 10 (1 -1 - 2 - 3 - 3) setae; endopod of 5 segments, with 2/2/1/2/5 (4 -1) setae; exopod with 4 natatory setae. Maxilliped 2 ( Fig. 2G): basis with 4 setae; endopod of 2 segments bearing 0/5 setae; exopod as for maxilliped 1. Maxilliped and pereiopods 1-4 present as unsegmented buds. Abdomen ( Fig. 2H) of 5 segments plus telson. Segments 2 and 3 with small dorsolateral protuber­ ances, those of segment 2 joined by a slightly raised dorsal ridge. Segments 2-5 each with a pair of short setae dorsally, near posterior margin of segment. Segments 4 and 5 widening posteriorly to a trilobed, spatulate telson. Median lobe of telson projecting beyond lateral lobes, which are posterolaterally serrate and terminate in a stout, minutely serrate spine with a further small spine laterally; between median lobe and each lateral lobe are 3 setose processes. Chromatophore pattern ( Fig. 2A,H; Table 1 ): lateral carapacial chromatophore well developed, with a single centre; abdominal segments 2-5 (Fig. 2H) with paired yellow and black chromatophores, the latter coalesced in segments 2-4; black subintestinal chromatophore of telson proximal to yellow lateral intestinal chromatophore. Material examined. AUTIIOR'S PERSONAL COLLEcnON. Waitemata Harbour, from Perna canaliculus: Westmere reef, 15 Oct 1981, 15 Dec 1981, 10 Feb 1982, 26 Apr 1982, 26 May 1982, numerous specimens; Stanley Point, 11 Feb 1981, 199; Narrow Neck, 11 Apr 1982, 49, 2g 9, 26. Harris Bay, Banks Peninsula, from Mytilus edulis aoteanus and P. canaliculus , 28 Jan 1982, 69, Ig9, 36. Camp Bay, Lyttelton Harbour, from M. e. aoteanus and P. canaliculus , 30 Jan 1982, 29, 5g 9, 20. AUCKLAND INSTITUTE AND MUSEUM. Kerikeri Inlet, Bay of Islands, from M. edulis , 30 Jan 1973, A. B. Stephenson, 21 '?, 21g'?, 90 ( AIM. 3924, 3925). Kaipara Harbour, from M. edulis , 17 Jan 1978, T. J. Bayliss, I'?, Ig'? ( AIM 3931). The Noises islands, from P. canaliculus , 23 Jan 1967, 25 Feb 1967, 25 Mar 1967, A. B. Stephenson, 2'?, 18g 9, 40 ( AIM. 3916--20). SE Ponui I., 5 Jun 1967, A. B. Stephenson, 20 ( AIM. 3922). Motuihi Channel, between Emu Pt and Home Bay, 8-10 fm, from P. canaliculus , 28 Mar 1930, A. W. B. Powell, Ig'? ( AIM. 3915). B.a.2, suction dredge off Devonport Wharf, Oct 1927, Mrs Movat, Ig'? (det. E. W. Bennett; AIM. 3914).?Manukau Harbour, Ig'? (det. E. W. Bennett; AIM. 3926). Matakana I., Tauranga, 4 ft, assoc. with Mytilus , 30 Jan 1972, A. B. Stephenson, 16 ( AIM. 3923).

NATIONAL MUSEUM OF NEW ZEALAND. Bay of Islands, dredged, Dec 1956, J. C. Yaldwyn, 19 (det. M. Scott, 1959; Cr. 949). Coromandel, from Chione stutchburyi , 31 Dec 1964, D. P. Orwin, 2g 9 (Cr. 2530). Long Bay, Auckland, from P. canaliculus, Feb 1950 , R. K. Dell, 69, 11g 9, 80 (Cr. 942). Waiheke 1., G. Chamberlain, 59,18 g'?, 40 (det. M. Scott, 1959; Cr. 944, 953). Off Ponui 1., Hauraki Gulf (36°51'S, 175°15'E), 10--15 fm, 29 Aug 1956, J. C. Yaldwyn, Ig9 (det. M. Scott, 1959; Cr. 945). Levin, from mantle cavity of toheroa: Waitarere Beach, 8 Nov 1951, 199 (det. M. Scott, 1959; Cr. 948); Hokio Beach, 5 Aug 1956, F. D. W., 19 (det. M. Scott, 1959; Cr. 946). Otaki Beach, from toheroa, Paphies (Mesodesma) ventricosa , 16 Sep 1978, G. H. Brandeis, 199 (Cr. 2527). Wellington Harbour: N end of Somes 1. from 48 Perna canaliculus , 27 Jul 1964, P. H. J. Castle, 29, 7g 9, 10 (Cr. 1496); Somes 1., from P. canaliculus , 8 Apr 1957, J. Moreland, 179, 6 g'?, 26 (Z. Cr. 676); off Days Bay, 4 fm, among algae, 19 Jan 1953, R. K. Dell and J. M. Moreland, 10 (Cr. 2525). Days Bay, from P. canaliculus , 4 Apr 1954, R. A. Falla, 19 (Cr. 954); wharf at light [sic], 16 Feb 1957, R. K. Dell, 69, 10 (Cr. 958); wharf piles, 12 Apr 1957, P. M. Ralph, 19, 199 (det. M. Scott, 1959; Cr. 950); Kau Pt, from 30 mussels, 20 Aug 1962, P. Castle, 2'?, l2g'?, 20 (Cr. 2517); Kau Pt, from 43 P. canaliculus, 10 Feb 1962, J. Moreland, 49, 8g 9 (Cr. 1197); Kau Pt, from P. canaliculus , 19 May 1962, J. Moreland, 16 (Cr. 1198); Karakara [sic] Bay, from P. canaliculus , 16 Feb 1948, R. K. Dell, 4g'? (det. M. Scott, 1959; Cr. 943). Totaranui, Nelson, free-living in eorallina, May 1963, R. K. Dell, 10' (Cr. 1313). Long 1., Queen Charlotte Sound, from M. edulis , 29 Sep 1963, M. A. Crozier, 19 (Cr. 2526). Tory Channel, Marlborough, 1957, M. Kirk, 199 (Cr. 957). Ketu Bay, Pelorus, mantle cavity of mussels, 1 Jan 1956, 19, 199, 10 (det. M. Scott, 1959; Cr. 947).

J. B. JONES PERSONAL COLLEcnON. Specimens from Ahipara, Wellington area (Ward 1., Seatoun beach, Eastbourne), and Marlborough Sounds (East Entry, Pelorus Sound, Kenepuru Sound) collected from P. canaliculus and M. edulis . Kaipara Harbour, from raft 1 m deep, Pacific oyster, 19. From Crassostrea , 10. Waitangi, from 16 Chione , 1 Aug 1973, 2g 9, 10.

CANTERBURY MUSEUM. Ponui passage, Aug 1915, W. J. Barr, 40 (Pinnatheres schauinslandi det. E. W. Bennett, 18 Nov 1930; AQ 2388), 19, 7g 9, 10 (det. E. W. Bennett, 18 Nov 1930; AQ 2250). Kaikoura, Jan 1936, H. R. Millar, 39, 199 ( AQ 2279). New Brighton, in polyzoan FluslTella binderi (drift from Sumner), 2 Jul1927, E. W. Bennett, 10 ( P. schauinslandi det, E. W. Bennett, 18 Nov 1930; AQ 2384). New Brighton, Hutton, 5'? (det. E. W. Bennett, 18 Nov 1930; AQ 2389). Sumner, C. Wood, 9'? ( AQ 2270). Robin Hood Bay, 30 Aug 1913, R. Bigg-Wither, 3d ( P. schauinslandi det. E. W. Bennett, 18 Nov 1930; AQ 2380). Riverton, 10 ( P. schauinslandi det. E. W. Bennett, 18 Nov 1930; AQ 2371, ex Chilton Collection), 19 {det. E. W. Bennett. 18 Nov 1930; AQ 2367, ex Chilton

.

Collection). Stewart I., in Mytilus, Apr 1916 , W. Traill, 10 ( P. schauinslandi det. E. W. Bennett, 18 Nov 1930; AQ 2304), 17!?, 20 (det. E. W. Bennett, 18 Nov 1930; AQ 2525). Chatham Island, Hutton, 5!? (det. E. W. Bennett, 18 Nov 1930; AQ 2392).

UNIVERSITY OF CANTERBURY. Camp Bay , Lyttelton Harbour, from Perna canaliculus, 1981 , A. S. Baxter, 4!?, 50.

Distribution. North and South islands, Stewart Island, and the Chatham Islands. Intertidal to 30 m. Endemic. The record of this species from Bass Strait, Tasmania ( Rathbun 1923, Guiler 1952) has been referred by Pregenzer (1979) to Pinnotheres hickmani (Guiler},

Hosts. Known hosts are Perna canaliculus , Mytilus edulis aoteanus , Aulacomya ater maoriana (Iredale), Oassostrea gigas (Thumberg), Paphies ventricosa (Gray) , and Chione stutchburyi (Gray) . Reported incidence in P. canaliculus ranges from zero to 70.27% ( Jones 1978, Hickman 1978, Baxter 1981). Stead (1971, p. 9) noted that several of the P. canaliculus examined during a survey of Pelorus Sound mussels contained a pea crab, and that pea crabs were also found occasionally in M. e. aoteanus. Baxter (1981) found 1.96-2.30% of A. a. maoriana from the Avon-Heathcote Estuary and Lyttelton Harbour to contain Pinnotheres novaezelandiae . In the C. stutchburyi populations studied by Larcombe (1971) the incidence of pea crabs, probably this species, was less than 1 %.

Hard-stage males and females have been taken free-living in Corallina , in subtidal algae, and on wharf piles.

Remarks. Filhol (1885a, b) described this species from specimens collected from Mytilus edulis at Golden Bay ("Baie du Massacre"). The syntypes (Museum National d'Histoire Naturelle, B. 8323) are 2 hard-stage females-described by Filhol as male-in excellent condition, and 11 mature females (2 gravid), most of which are badly damaged. The largest mature female (carapace width 14.9 mm, without abdomen, left chela detached) is referable to the new species of Pinnotheres described below, not to P. novaezelandiae as understood by recent authors. This suggests that Filhol's material came from more than one locality and host. Of the remaining syntypes, 3 are readily determinable as P. novaezelandiae and 7 are badly damaged, making identification difficult. I have chosen here to select the larger hard-stage female (carapace width 9.1 mm) as the lectotype of P. novaezelandiae and to designate the remaining syntypes, with the exception of the largest mature female mentioned above, paralectotypes.

Lenz (1901) examined one of Filhol's specimens, together with material from Mytilus from French Pass, and noted several inaccuracies in Filhol's figures (see also Scott 1961). In addition Lenz described a new species, Pinnotheres schauinslandi , based on 2 specimens of unstated sex collected with P. novaezelandiae from the same host at the same locality.

Chilton (1911) suggested that Lenz had in fact described the male (i.e., hard-stage) of P. novaezelandiae as a new species, a view that Scott (1961) supported. Only Bennett (1964) has recognised P. schauinslandi as a valid species. Examination of Canterbury Museum material determined by Bennett in 1930 shows that he assigned all hardstage Pinnotheres to P. schauinslandi and all softstages to P. novaezelandiae . Apparently Bennett was unaware of the complex polymorphic development of the Pinnotheridae , and regarded hardand soft-stage pea crabs as different species rather than developmental morphs. There are small differences between hard-stage P. novaezelandiae and Lenz's description of P. schauinslandi (see Bennett 1964, p. 79; Jones 1977), and until Lenz's material (if extant) is examined the status of P. schauinslandi will remain open to doubt. However, given the original description, the host, and the circumstances of collection I agree with Chilton (1911) and Scott (1961), and include P. schauinslandi in the synonymy of P. novaezelandiae .

In a postscript, Scott (1961) stated that the pea crab recorded by Dell (1960) as Pinnotheres sp. from Nemocardium pulchellum Gray in Petre Bay, Chatham Islands, was a hard-stage female, not a male as Dell stated, and that it differed from P. novaezelandiae in possessing a row of setae on the dorsal margin of the dactylus of the chelae. In fact, most females of P. novaezelandiae also possess these setae. I could not find Dell's specimen in either the Canterbury Museum or the National Museum, where the 1954 Chatham Islands Expedition material was deposited, so the identity of this specimen remains uncertain.

The persistent, erroneous records of the European Pinnotheres pisum (L.) have been discussed by Scott (1961) and Bennett (1964), and Pregenzer (1979) has compared the adults of P. pisum and P. novaezelandiae in some detail. The differences in the first-stage zoeae of the 2 species are briefly discussed by all 3 authors. P. novaezelandiae has also been confused with the Australian P. hickmani (Guiler) ; Pregenzer (1979) has distinguished these 2 species.

Since Jones's (1978) study of the natural history of P. novaezelandiae the incidence of this species in natural and cultivated populations of P. canaliculus has been discussed by Hickman (1978), and Baxter (1981, 1982) has studied aspects of its ecology and physiology.

Tables

Table 1 Chromatophore patterns of first-stage zoea larvae of Pinnotheres novaezelandiae and P. atrinicola (-, chromatophore absent).

		novaezelandiae	atrinicola
PRIMARY SYSTEM
NEURAL GROUP
1.	Supracerebral
2.	Antennular
3.	Antennal	dichromatic,	black
		black/yellow
4.	Labral
5.	Mandibular	black	black
6.	Maxillary	black	black
7.	Maxillipedal	black	black
8.	Lateral
	intestinal	yellow	yellow
9.	Subintestinal	black, paired	black, paired
		in segment 5	in all segments
		and telson
VISCERAL GROUP
10.	Median gastric	black	black
11.	Precardiac	black	black
12.	Subcardiac	dichromatic,	dichromatic,
		black/yellow	black/yellow
13.	Postcardiac
SECONDARY SYSTEM
14.	Posterior
	carapacial
15.	Posteroventral	dichromatic,	dichromatic,
		black/yellow	black/yellow
16.	Lateral	dichromatic,	dichromatic,
	carapacial	black/yellow	black/yellow
17.	Dorsal carapace
	spine
18.	Maxillipedal
19.	Optic
20.	Median ocular	black	black
	centre