Callipallenidae Hilton, 1942

Staples, David A., 2014, A revision of the callipallenid genus Pseudopallene Wilson, 1878 (Pycnogonida, Callipallenidae), Zootaxa 3765 (4), pp. 339-359 : 342-346

Family Callipallenidae Hilton, 1942

Synonymies

Pallenidae Sars, 1891: 31 .— Barnard, 1954: 105 (for most of the earlier literature). Callipallenidae Hilton, 1942b: 281 .— Stock, 1954: 28.

Diagnosis. Octopodus. Body oval or elongate with lateral processes distinctly separated or touching. Abdomen fusiform. Ocular tubercle on mid to posterior half of cephalon. Eyes four or eight. Proboscis directed obliquely downwards or carried in line with the trunk, distally mamilliform, truncate or tubiform, jaws triradiate. Chelifores well-developed, directed obliquely downwards or carried in line with the trunk, chelifore scapes one-segmented, sexual dimorphism variable, chelae large, carried transversely in front of the mouth or in line with the chelifore scape, fingers toothed, smooth, rugose or with marginal lobe. Ovigers 10-segmented in both sexes or sixsegmented in some males, fifth segment of male oviger longer than that of female and with distal apophysis; strigilis and terminal claw present or absent, spines compound or simple, in single row. Tarsus much shorter than propodus. Genital pores on ventral surface of second coxa, in males on legs 3 and 4, in females on all legs.

The high level of morphological variability amongst callipallenid genera places into question their natural grouping within the family Callipallenidae . Whilst auxiliary claws are not a good taxonomic character, other characters such as femoral cement glands and palps are regarded by this author to be of significance. The variable presence of auxiliary claws in the genera Nymphon , Pycnogonum and Ammothella is well documented and accepted. In the family Callipallenidae auxiliary claws are present in all species within two genera and are variably present in two others. About twice as many genera are diagnosed with femoral cement glands as those that have not, and in one genus they are variably present. One to four segmented palps are found in ten genera and may be present in males only, or in both sexes.

Based on this analysis, species presently assigned to Pseudopallene fall into three primary groups herein assigned to independent genera.

Genus Pseudopallene Wilson, 1878 ( Figure 2 F)

Pseudopallene Barnard 1954: 107 for earlier synonyms. Pseudopallene Stock, 1954: 61 .

Diagnosis. Trunk segmented, smooth or spinous, neck region of cephalon broad, anterior region of cephalon swollen; lateral processes longer than basal width, narrowly separated. Abdomen typically short, rounded distally. Four eyes, usually pigmented. Ocular tubercle prominent, on posterior half of cephalon. Proboscis tip with dense fringe or tuft of long setae. Palps one-segmented buds in both sexes. Chelifores carried in line with trunk, chelae palm swollen, fingers robust, without teeth but with a node variably present. Oviger claw long, pointed with one or more distinct sharp teeth usually confined to inner margin. Legs smooth or spinous, auxiliary claws absent. Cement glands not present.

Type species of the genus: Pseudopallene circularis ( Goodsir, 1842) .

Remarks. Three of the six species assigned here to Pseudopallene have been re-examined and can be assigned to the genus with confidence. These are P. circularis ( Goodsir, 1842) , P. spinipes ( Fabricius, 1780) and P. brevicollis ( Sars, 1888) . One-segmented palp buds are present in each species. These three species are recorded from the north Atlantic. The oviger spines of P. spinipes and P. brevicollis are compound whereas those of P. circularis are simple. Three other species have not been seen in this study and are assigned to Pseudopallene pending reexamination of the types or the availability of additional material. Palp buds were not mentioned in the descriptions of these species but history has shown that these buds have been overlooked owing to their tiny size and their ventral (rather than lateral) position on the cephalon. Palp buds are presumed to be present in these species. Two species, Pseudopallene collaris Turpaeva, 2002 and P. pacifica Losina-Losinsky, 1961 are both recorded from the Kuril Chain of islands in the Sea of Okhotsk northwestern Pacific and are biogeographically consistent with other records in the northern hemisphere. The third species P. centrotus Pushkin, 1990 is recorded from South Georgia (54° 53' S, 34° 30' W). This is the only species not recorded from the Boreal-Arctic zone. Despite the geographic disparity, P. centrotus is much like northern hemisphere species of Pseudopallene with which it agrees generally in habitus and in the shape of the oviger claw. As with P. collaris the proboscis is orientated ventrally. Pushkin did not record the type of oviger spine. Pushkin distinguished P. centrotus from its congeners primarily by the possession of multiple spines on the lateral processes and legs. Turpaeva distinguished P. c o l l a r i s from P. circularis largely by possession of a prominent ‘collar’ around the base of the proboscis and the ventral orientation of the proboscis. Pseudopallene collaris shares possession of smooth oviger spines with P. circularis . Unfortunately Losina-Losinsky did not describe the oviger spines and distal claw of P. pacifica in detail other than to observe that there are few spines on the distal segments. The shape of the chelifore fingers and long abdomen are inconsistent with its congeners and should it be determined that palp buds are not present then the status of this species will need to be reassessed. Based on material loaned by the Smithsonian Institution National Museum of Natural History, the opportunity is taken here to update the early descriptions of the three species examined and to provide additional figures and comments. I am grateful for one additional specimen of P. circularis provided by Dr Roger Bamber. This specimen has been registered and lodged with Museum Victoria.

Distribution. North Atlantic; N-W Pacific; South Georgia, Antarctica.

Pseudopallene circularis ( Goodsir, 1842) Type species of the genus by designation. Figures 1 C–D

Cordylochele microspines Hilton, 1942c: 39 –40

Cordylochele setospines Hilton, 1942c: 39

Pseudopallene setoso Hilton, 1942c: 39

Pseudopallene spinosa Hilton, 1942c: 39

Pseudopallene circularis . Hedgpeth, 1948: 206 (for most other early synonymies).— Stock, 1953a: 41 — Stock, 1953b: 297 — Hedgpeth, 1963:1333.

Material examined. USNM 153832. USNM Acc. No. 188318. Ovigerous male. Canada, Labrador, off Lowland Point, 53° 50'N, 059° 25'W. Blue Dolphin stn 50-5. Coll. D. C. Nutt, Dredge, depth 20fms. 25 Jul 1950. Det. C. A. Child. NMV J46797 Ovigerous male. Off Northumberland, UK, ‘The Trink’, depth 30–40 m, 20 Mar 1989. Det. R. N. Bamber.

Description of male. (Specimen USNM153832).Trunk compact, ovate in outline, second segment with two side-by-side mid-dorsal, spine-tipped tubercles, segment 3 with two spine-tipped tubercles placed one behind the other. Neck very short, crop strongly expanded, three strong spine-tipped tubercles present at base of each chelifore scape insertion. Lateral processes almost touching, distally fringed with six to eight spine-tipped tubercles, suture line separating process from the trunk not present. Abdomen length four times greatest width. Eyes pigmented, width of ocular tubercle at base a little greater than height, truncate dorsally with single mid-dorsal papillae. Proboscis swollen proximally, narrowing in mid-region before dilating again and terminating in a mamilliform tip. A dense fringe of long setae lines the jaws. Palp buds claviform, each palp positioned at base of proboscis margins ( Fig 1 D). Chela immoveable finger with a prominent node at about half its length, both fingers with heavily sclerotized margins; tips of fingers do not meet, moveable finger rests on outer margin of immoveable finger, proximal margin of chelifore scape much narrower than remainder of scape. Oviger base inserted beneath anterior margins of first lateral processes, segment 5 with seta-tipped distal apophysis, segments 7–10 lined with simple, curved spines according to the formula 4:4:4:3 (specimen NMV J46797 ); spines almost as long as width of corresponding segment, terminal claw much shorter than segment 10, with single tooth on inner and outer surfaces. Legs covered in spine-tipped tubercles, the terminal spine is longer than basal tubercle which is covered in tiny setae; femur with two, smooth, rounded protuberances on ventral margin, a similar single protuberance is present on second tibia. Propodal heel not defined, heel region and sole lined with five to six strong spines gradually reducing in size distally, distal two spines very much smaller. Genital pores small, inconspicuous.

Remarks. Hedgpeth (1963) was the first to observe that all of Hilton’s proposed species are various growth stages of P. circularis . Sars (1891) described the oviger spines as ‘irregularly disposed’ but in the specimen examined they are in a single row. One or two strong distal setae on segments 7–10 potentially make the spine count ambiguous. The rounded protuberances noted here on the ventral margin of the femur and second tibia are recorded by Sars (1891) on the femur and both tibiae. The description of specimens being ‘covered in hairs’ by Goodsir (1842) and ‘hairy’ by Wilson (1878b) may be attributed to terminology but could also indicate that more than one species has been assigned to P. circularis . In any event these descriptions are not consistent with the specimens examined.

Distribution. Boreal-Arctic, widely distributed in colder waters of the North Atlantic (Okhotsk Sea to West Greenland, south to the Firth of Forth, southern Norway, Cape Cod).

Type locality Firth of Forth, Scotland

Pseudopallene brevicollis ( Sars, 1888) Figures 1 E, F

Cordylochele brevicollis Sars, 1888 .— Sars, 1891: 51. Pseudopallene brevicollis Stock, 1953b: 297 .

Material examined. USNM 128655, 2 females. Canada, Labrador, Blue Dolphin stn. BD-43, near Goose Bay, 53° 52'N 059° 19'W, Otter Trawl, 30–35 fms, 11 Jul 1950, Det. C. A. Child, Dec. 1969.

Description of female. Trunk, smooth, segmented. Proboscis and chelifores lying almost in a horizontal plane, proboscis and chelifores only slightly down-curved. Neck region of cephalon well defined, about as long as wide. Lateral processes about as long as wide, narrowly separated. Abdomen short, curved downward. Ocular tubercle a low mound, hardly raised, eyes darkly pigmented. Proboscis narrowing in mid-region, widening to maximum width before terminating in obtuse frontal region, lips slightly extended with dense tuft of long setae ( Fig. 1 A). Chelifore scapes about as long as proboscis, smooth, without notable proximal constriction, fingers pointed, with proximal ‘fleshy’ spinous pads on their ventral surfaces ( Fig 1 F). Palp buds curved inward, each positioned between the outer margin of the proboscis and the inner margin of the chelifore scapes. Oviger bases occupying mid-region of neck, segment 4 with strong proximal swelling on outer margin, strigilis spines strongly curved inwards, broad and with indistinct crenulate edges, spine formula; 12:12:11:12, claw about half length of segment 10, with seven teeth along inner margin (three distal teeth blunt, four proximal teeth pointed). Legs covered in short setae, propodus, slightly curved, length> three-times length of tarsus, heel only slightly raised, two large spines, distal-most spine longest, terminal claw long, a little shorter than length of propodus. Genital pores large, bulging.

Remarks. The smooth trunk and narrower neck region of the cephalon set this species apart from its congeners. The fleshy pads at the base of the moveable finger are reminiscent of Pallenopsis Wilson, 1881 but from which it otherwise differs considerably. Sars, (1891) observed the two small, tubiform palp buds.

Distribution. Arctic, sub Arctic, North Atlantic.

Pseudopallene spinipes ( O. Fabricius, 1780) Figures 1 G–H

Pseudopallene spinipes Sars, 1891: 42 (for earlier synonymies).

Phoxichilus spinipes Norman, 1908: 207 .

Pseudopallene spinipes Stephensen, 1913b: 387 . — Schimkewitsch 1930. — Stephensen 1933a: 21.— Stephensen 1933c: 36 – 38.— Hedgpeth, 1943: 88.— Stock 1953b: 297.

Material examined. USNM 80761, 1 female. Canada, Baffin Bay; east of Cobourg I., 75° 40'N 078° 40'W, 140– 210 ft. Bartlett R. A., 3 Aug 1935, Det. Hedgpeth, J. W.

Description of female. Trunk compact, segmented, mid-dorsal region smooth. Neck very short, strongly expanded to form a well-developed cowl at the base of each chelifore scape insertion, four-five smooth, pointed tubercles on anterior margins of cowls. Lateral processes separated by about half their basal width, each bearing numerous spinules and with a dorsolateral process on the anterior and posterior margins, fourth lateral process with dorsodistal median spine, suture line separating trunk and lateral processes not present. Abdomen length about three times greatest width, with several spine-tipped processes. Four eyes. Ocular tubercle about as high as wide, two low, dorsal papillae. Proboscis carried horizontally forward of the cephalon, swollen proximally, narrowing slightly in mid region before dilating again and terminating in a mamilliform tip, with fringe of long setae. Chelifore scape with numerous pointed processes, largest on distal mid-dorsal margin, flanked by two smaller spines, proximal constriction of scape not evident. Chelae finger margins heavily sclerotized, tips meeting, proximal to the tips there is a single node on the moveable finger which interlocks with two corresponding side-byside nodes on the immoveable finger. Dorsal surface of moveable finger and surface of palm covered in spinules, palm much longer than fingers, two palp buds present on the ventrodistal surface of cephalon, set wide-apart in line with the outer margins of the cephalon ( Fig. 1 H). Oviger bases inserted beneath anterior margin of first lateral processes, segment 5 with prominent distal apophysis, segments 7–10 lined with compound spines according to the formula 6:5:4:6., terminal claw pointed, about equal in length to segment 10, with three sharp teeth on inner margin. Legs covered in spine-tipped processes, basal part smooth, terminal spine longer than basal process. Propodal heel present, with two or three large spines followed by about eight smaller spines on sole, tarsus with one large spine. Genital pores not evident although the specimen otherwise appears to be a mature adult.

Distribution. Boreal-Arctic, widely distributed: Greenland, Baffin Bay, Western Norway, Kara Sea, Franz Joseph Land.

Figures

FIGURE 1. Pseudopallene brevicollis; male, (USNM 128655), A, detail of proboscis tuft. Meridinale watsonae; B, detail of proboscis fringe. Pseudopallene circularis; male, (USNM 153832), C, trunk dorsal view, D, male, (NMV J 46797), palps ventral view. Pseudopallene brevicollis; E, female, (USNM 128655), trunk dorsal view; F, ventral view of palps. P seudopallene spinipes; G, female. (USNM 80761), trunk, dorsal view; H, ventral view of palps.

FIGURE 2. Anoplodactylus evansi; A, southern Australian specimen; B, Tweed Heads, northern N. S. W. specimen. Pallenopsis macneilli; C, specimen from Eaglehawk Neck, Tasmania. Photo Andrew Newton; D, specimen from Tollgate Is. NSW. Photo Andrew Green. Meridionale chevron; male holotype, (SAM E 3681), E, anterior view. Meridionale sp.; male, (NMV J 57280), F, lateral view of typical form.