Ischyrocerini Kroyer 1838
Tribe Ischyrocerini Krøyer, 1838 Supplementary Table S2 Type genus. Ischyrocerus Krøyer, 1838 Diagnosis (with changes from Just (2017) in bold). Antennae : slender, antenna 1 with accessory flagellum (occasionally vestigial). Mandible : palp with 3 articles, the third expanded distally (occasionally...
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| Summary: | Tribe Ischyrocerini Krøyer, 1838 Supplementary Table S2 Type genus. Ischyrocerus Krøyer, 1838 Diagnosis (with changes from Just (2017) in bold). Antennae : slender, antenna 1 with accessory flagellum (occasionally vestigial). Mandible : palp with 3 articles, the third expanded distally (occasionally similar in shape to the second) . Coxae 1–4 : progressively deepening, subrectangular to oval (occasionally coxa 1 much smaller than and mostly obscured by coxa 2 and differing in shape), margins entire. Gnathopod 1 : carpus shorter than the propodus (occasionally longer), propodus oval to weakly subchelate. Gnathopod 2 : propodus in adult male ( and occasionally in the female ) moderately to strongly enlarged compared to gnathopod 1, of varying shape ( occasionally hardly modified ). Pereopods 3–4 : merus moderately to fully overlapping the carpus anteriorly, dactyl shorter than the carpus (occasionally longer ). Pereopods 5–7 : of similar form, increasing in length backwards ( occasionally 6 larger than 5 and 7 ). Urosomites : 1–3 free. Uropods 1 and 2 : peduncle without distoventral corona of spines , with 2 subequal rami with or without an underlying peduncular spinous process (occasionally uropod 2 outer ramus modified ). Uropod 3 : peduncle long ( occasionally short ), broad proximally, narrow distally, biramous ( occasionally uniramous), outer ramus terminating in cusps and/or spine(s ). Telson : entire, with one to many dorsally or apically projecting setae or spines . Component genera. Jassa Leach, 1814; Ischyrocerus Krøyer, 1838; Paradryope Stebbing, 1888; Microjassa Stebbing, 1899; Parajassa Stebbing, 1899; Hemijassa Walker, 1907 Isaeopsis K.H. Barnard, 1916; Pseudischyrocerus Schellenberg, 1931; Bathyphotis Stephensen, 1944; Ventojassa J.L. Barnard, 1970; Neoischyrocerus Conlan, 1995; Scutischyrocerus Myers, 1995; Ruffojassa Vader & Myers, 1996; Veronajassa Vader & Myers, 1996; Alatajassa Conlan, 2007; Myersius Souza-Filho & Serejo, 2014; Pleojassa n. gen.; Plumulojassa n. gen. Changes to Ischyrocerus and Neoischyrocerus Ischyrocerus is primarily a cold water, Northern Hemisphere genus, captured from deep trawls (Stephensen 1944; Gurjanova 1951) as far north as the high Arctic, but also found in the intertidal and shallow subtidal zone (J.L. Barnard 1962). It has been extensively found in the Southern Hemisphere as well, though mostly in warmer waters (Myers 1995, 1997; Just 2009). This large genus requires revision and may prove to be less cosmopolitan than previously thought by J.L. Barnard & Karaman (1991). Three genera have been created for warm water Ischyrocerus -like species: Neoischyrocerus Conlan, 1995 (4 species), Coxischyrocerus Just, 2009 (2 species) and Tropischyrocerus Just, 2009 (2 species). These genera embrace species in which the male develops an enormously lengthened and pendulous gnathopod 2 (about 200–300% the length of gnathopod 1) with an anteriorly rounded ischium and a very long propodus with the palm nearly the full length of the propodus, and the proximal end of the palm marked by a bulge next to the carpus (the S. Californian N. claustris (J.L. Barnard, 1969), N. chinipa (J.L. Barnard, 1979) from the Galapagos and Pacific Panama, N. vidali Ortiz & Lalana, 2002 from the Cuban Caribbean, C. inexpectatus (Ruffo, 1959) from the Mediterranean Sea and T. socia (Myers, 1989) from Bora Bora), a tooth-like projection (e.g., N. lilipuna (J.L. Barnard, 1970) from Hawaii and T. pugilus Just, 2009 from Australia), or with neither (e.g., C. rhombocoxus Just, 2009 from Australia). The dactyl may be the full length of the propodus or shorter, the length growth related. By comparison, the female’s gnathopods are similarly sized with the second only slightly larger than the first. Other commonalities are antennae with long filtering setae that are not pediform or sexually dimorphic, a 2-articulate accessory flagellum with the second article minute, pereopods 3 and 4 with little overlap of the merus over the carpus, a well developed peduncular spinous process under the rami of uropod 1, a spiny peduncle of uropod 3 with the outer ramus bearing a row of minute cusps and the inner ramus tipped by a small spine, and the telson with a pair of strong, dorsally projecting spines. The difficulty with these genera is where species cross the generic boundaries. Examples are: enlarged coxa 2 relative to coxa 1 in the adult male ( C. rhombocoxus and N. claustris ), similar gnathopod propodus appearance as noted above, and similar female gnathopod palms (convex in all species in the three genera except for T. pugilus ), pereopod 3 and 4 propodus posteriorly spinose ( N. lilipuna , N. vidali , T. socia and C. inexpectatus) . The generic-level differences among the genera therefore recede into issues of sexual variation (e.g., enlargement of coxa 2 relative to coxa 1 that was used to define Coxischyrocerus but also occurs in Neoischyrocerus , or the modified pereopod 5 basis shape in adult males of C. rhombocoxus and C. inexpectatus ). Therefore, Coxischyrocerus Just, 2009 and Tropischyrocerus Just, 2009 are herein merged into the senior genus Neoischyrocerus Conlan, 1995. Myers (1995, 1997) noted the need for diminutive Indo-Pacific species placed at that time in Ischyrocerus or Jassa to be placed in their own genus and these are also included, as noted below. Genus Neoischyrocerus n. comb. Supplementary Table S3 Type species. N. claustris J.L. Barnard, 1969 Diagnosis. (with differences from Ischyrocerus in bold). Body length at maturity 1–2 mm (usually), rarely 4–6 mm. Tropical and warm temperate distribution, collected from algae, sponges, corals or from a spiny lobster, 0–16 m, 9– 40°N and 5– 34°S. Pereon : dorsally smooth (most species), ridged or carinate (some species). Antenna 1 : accessory flagellum 2 articles (second minute), projecting forward or flush with the flagellum; antennae 1 and 2 peduncles subequal in width or antenna 2, 10% wider (based on comparison of antenna 1 peduncle article 2 with antenna 2 peduncle article 4), peduncular setae and setal pattern similar to antenna 1, or slightly shorter, not plumose. Gnathopod 2, adult male : 190–350% the length of gnathopod 1, basis and propodus especially elongate coxa 1, 60–110% the depth of coxa 2; basis concave or sinuous ischium anteriorly rounded; propodus often slender (posterior length 180–400% of central width), palm nearly the full length of the propodus, often with a bulge or tooth defining it proximally at the junction of the carpus (sometimes palm continuous with the carpus), sometimes centrally toothed instead and with shallow bulge or teeth at the junction of the dactyl dactyl half or nearly the full length of the propodus. Gnathopod 2, juvenile male : shorter than the adult male gnathopod 2, palm bearing 1–2 strong spines about midway along the length of the propodus, dactyl ending at the spines. Gnathopod 2, female : only slightly larger than gnathopod 1, palm of the propodus convex (usually) or shallowly concave (rarely). Pereopods 3 and 4 : propodus, posterior margin bearing spines or setae coxa 4, posterior margin straight, not shallowly concave. Pereopod 5, male : similar to but shorter than pereopods 6 and 7 or variously modified with posteriorly concave basis or posteriorly expanded and spinose merus. Uropod 1 : peduncle with short ventrodistal spinous process underlying the rami, length ~15–35% of the outer ramus length. Uropod 3 : peduncle bearing 1–2 rows of spines dorsally, ending in a single spine at the distal margin, but without a corona of spines around the margin or setae; rami without spines mid-dorsally , outer ramus subequal to or shorter than the inner and bearing 3–8 minute dorsal cusps apically, without (rarely with) a small apical straight spine . Component species (with transferred species in bold). Ischyrocerus longimanus (Haswell, 1879) (Australia); I. parvus Stout, 1913 (California); I. carinatus K.H. Barnard, 1916 (South Africa); I. gorgoniae K.H. Barnard, 1940 (South Africa); I. ctenophorus Schellenberg, 1953 (South Africa); Coxischyrocerus inexpectatus (Ruffo, 1959) (Mediterranean, Red Sea?); N. claustris J.L. Barnard, 1969 (California); N. lilipuna J.L. Barnard, 1970 (Hawaii); I. oahu J.L. Barnard, 1970 (Hawaii); I. oahu oahu J.L. Barnard, 1970 (Hawaii); N. chinipa J.L. Barnard, 1979 (Galapagos Islands and Panama); I. oahu armatus Ledoyer, 1979 (Madagascar); Tropischyrocerus socia (Myers, 1989) (Bora Bora); I. mediodens Myers, 1995 (Papua New Guinea); I. parma Myers, 1995 (Papua New Guinea); I. apiensis Myers, 1997 (Samoa); N. vidali Ortiz & Lalana, 2002 (Cuba); C. rhombocoxus Just, 2009 (Australia); T. pugilus Just, 2009 (Australia). Remarks. Species of Ischyrocerus were transferred to Neoischyrocerus if they demonstrated at least one key character (grossly enlarged and pendulous male gnathopod 2 similar in shape to that of others in the genus; dactyls with comb-like striae as noted in J.L. Barnard (1970), Conlan (1995) and Ortiz & Lalana (2002); similar spination on uropod 3). Presence or absence of these striae were not mentioned by other authors, therefore questioning as to whether this character had been looked for. Mouthpart characteristics were not widely described, but may be useful for generic definition, especially the clavate vs slenderer shape of the mandibular palp and the presence/absence of a long apical seta on the maxilla 1 inner plate. Excluded but uncertain generic status . Ischyrocerus kapu J.L. Barnard, 1970 from Hawaii. The author based the generic assignment on a single male specimen. He noted its resemblance to N. lilipuna but also considered that it should be in a new genus. On balance, though, he assigned it to Ischyrocerus but noted that this was based on limited information because the specimen lacked antennae and pereopods and the female was also unknown. The male’s gnathopod 2 is unusual in having a long conical extension of the merus underneath the propodus, a feature that is not known for either Ischyrocerus or Neoischyrocerus . Myers (1995) stated that I. kapu is congeneric with other species being transferred to Neoischyrocerus . The male’s propodus is wider than in other members of Neoischyrocerus , but its uropod 3 resembles other species of Neoischyrocerus rather than Ischyrocerus . Further material demonstrating the species’ complete morphology is required before it can be confidently transferred to Neoischyrocerus or to a new genus. Genus Ischyrocerus n. comb. Supplementary Table S4 Type species. Ischyrocerus anguipes Krøyer, 1838 Diagnosis (with differences from Neoischyrocerus in bold). Body length at maturity 2–18 mm , though most species are> 5 mm long at adulthood. Primarily known from the Northern Hemisphere in cold temperate to polar waters from 32°N (La Jolla, California; J.L. Barnard 1969) to 81°N in the Arctic Ocean (Stephensen 1944), collected from algae, hydroids, crabs or substrate unknown, 1 to ~ 2000 m. Widely known in Europe but not in the Southern Hemisphere, although one species was found at 42°S off of Chile (J.L. Barnard 1964). Pereon : dorsally smooth (most species), ridged or carinate (some species). Antenna 1 : accessory flagellum 2 articles (second minute), flush with the flagellum or rarely projecting forward; antenna 2 peduncle article 4, 115–170% wider than antenna 1 peduncle article 2 (or rarely equal width), setae and setal pattern similar to antenna 1 or setae shorter, occasionally plumose. Gnathopod 2, adult male : 100–220% the length of gnathopod 1, basis usually not especially elongate, propodus elongate and slender or short and broad (rarely similar to gnathopod 1) coxa 1 70–140% the depth of coxa 2; basis concave or straight ischium anteriorly rounded or straight; propodus variably shaped, slender or broad (posterior length 120–290% of central width), palm only on the distal portion of the propodus or nearly the full length of the propodus, without a bulge or tooth defining it proximally at the junction of the carpus, palm variously toothed dactyl 33–75% the length of the propodus. Gnathopod 2, juvenile male : shorter than the adult male gnathopod 2, palm bearing 1–2 strong spines about midway along the length of the propodus, dactyl ending at the spines. Gnathopod 2, female : only slightly larger than gnathopod 1, palm of the propodus convex or concave. Pereopods 3 and 4 : propodus, posterior margin bearing setae but not spines; c oxa 4, posterior margin straight to shallowly concave. Pereopod 5, male : similar to but shorter than pereopods 6 and 7 or basis posteriorly concave; merus not differing. Uropod 1 : peduncle with short ventrodistal spinous process underlying the rami, length ~25–50% of the outer ramus length. Uropod 3 : peduncle bearing 0–2 rows of spines dorsally, ending in a corona of spines at the distal margin ( rarely a single seta or single spine) rami with 0–1 spines (outer), 0–4 spines (inner), outer ramus subequal to a third shorter than the inner and bearing 0–9 cusps, occasionally with 1–2 apical spines that are straight or dorsally recurved . Component species. Ischyrocerus anguipes Krøyer, 1838 (N. Europe and Arctic Ocean); I. latipes Krøyer, 1842 (Arctic Ocean); I. minutus Liljeborg, 1851 (N. Europe); I. megacheir (Boeck, 1871) (40°N– 80°N, Atlantic to Arctic Ocean); I. brevicornis (Sars, 1879) (E. Greenland, Arctic Ocean); I. tuberculatus (Hoek, 1882) (Barents Sea, 71°N– 77°N); I. tenuicornis (Sars, 1885) (N. Europe); I. nanoides (Hansen, 1887) (Arctic, Baffin Bay and W Greenland, 61°N – 81°N); I. megalops Sars, 1894 (N. Europe); I. commensalis Chevreux, 1900 (E. Atlantic Canada and Saguenay Fjord); I. brusilovi Gurjanova, 1933 (Russian waters); I. enigmaticus Gurjanova, 1934 (Kara Sea, 78°58ʹN); I. cristatus Gurjanova, 1938 (Sea of Japan); I. elongatus Gurjanova, 1938 (Sea of Japan); I. rhodomelae Gurjanova, 1938 (Sea of Japan); I. serratus Gurjanova, 1938 (Sea of Japan); I. hanseni Stephensen, 1944 (64°N, between Iceland and Greenland); I. albanovi Gurjanova, 1946 (Arctic Ocean); I. laptevi Gurjanova, 1946 (Arctic Ocean); I. chamissoi Gurjanova, 1951 (Russian waters); I. dezhnevi Gurjanova, 1951 (Russian waters); I. krascheninnikovi Gurjanova, 1951 (Russian waters); I. stephenseni Gurjanova, 1951 (Russian waters); I. pelagops J.L. Barnard, 1962 (California); I. hortator J.L. Barnard, 1964 (off Chile); I. malacus J.L. Barnard, 1964 (California); I. gurjanovae Kudrjaschov, 1975 (Kurile Islands); I. tzvetkovae Kudrjaschov, 1975 (Kurile Islands); I. fractus King & Holmes, 2004 (Ireland). Remarks. Species were retained in Ischyrocerus if they lacked the key characters noted above for Neoischyrocerus in the appearance of the male gnathopod 2, pereopod dactyls or uropod 3 spination pattern. Ischyrocerus fractus is the only species known in this genus where the male’s gnathopod 2 propodus is very little different from the female’s. It is also the smallest known at adulthood for this genus (2 mm). Excluded, but uncertain generic status. Ischyrocerus camptonyx Thurston, 1974b from subantarctic Signy Island is not Ischyrocerus . It is possibly an undescribed species of Jassa or synonymous with J. alonsoae , in which case Thurston’s name would take precedence. Jassa thurstoni Conlan, 1990 (called J. falcata form 2 by Thurston) and Pleojassa moorei n. sp. (called J. falcata form 3) are also known from Thurston’s collections there. For I. camptonyx , hallmarks of the genus Jassa, rather than Ischyrocerus are the spines at the tip of the antenna 2, the sinuous palmed gnathopod 2, the strong overlap of the merus over the carpus on pereopods 3 and 4, the typical Jassa -like uropod 3 with long peduncle lacking mid-dorsal spines (but with a corona of spines around the distal margin), a lateral setal brush and the strong hooked spines at the tip of the outer ramus, and the telson with a long seta at each corner rather than a spine. However, Thurston describes the uropod outer ramus “with three stout hooked spines dorsally near apex and a minute comb with five-six teeth laterally”, which does not correspond to his illustration and are not Jassa -like. Possibly, though, his description could be interpreted differently. One of the three spines may be the apically immersed, dorsally recurved spine typical of Jassa , the other two spines are cusps, and the five–six teeth are minute dorsal cusps proximal to the two large ones. If so, then this also speaks of I. camptonyx as being a Jassa , either its own species or synonymous with J. alonsoae . It is not a Pleojassa , even though the male’s second-gnathopod resembles that of P. moorei , because this genus lacks a gill on gnathopod 2 while I. camptonyx possesses one. Thurston considered that the few males available for study were juvenile because they all lacked a thumb as in Jassa . However, some of these specimens were larger than the adult, ovigerous females. The female allotype was 4.5 mm and the male holotype was 5.5 mm. This suggests that the male holotype is actually an adult that will not produce a thumb, in which case it is not Jassa . Therefore, until the range of variation can be assessed in Thurston’s specimens, this species should remain in Ischyrocerus with a question as to its proper generic placement. Uncertain status of : Published as part of Conlan, Kathleen E., 2021, New genera for species of Jassa Leach (Crustacea: Amphipoda) and their relationship to a revised Ischyrocerini, pp. 1-72 in Zootaxa 4921 (1) on pages 57-61, DOI: 10.11646/zootaxa.4921.1.1, http://zenodo.org/record/4496015 : {"references": ["Kroyer, H. (1838) Gronlands amfipoder beskrevne af Henrick Kroyer. Det Kongelige Danske Videnskabernes Selskabs Naturvidenskabelige og Mathematiske Afhandlinger, 7, 229 - 326. https: // doi. org / 10.5962 / bhl. title. 13747", "Just, J. (2017) A fresh look at the higher classification of the Siphonoecetini Just, 1983 (Crustacea, Amphipoda, Ischyroceridae) 12: with a key to all taxa. Zootaxa, 4320 (2), 321 - 338. https: // doi. org / 10.11646 / zootaxa. 4320.2.7", "Leach, W. E. (1814) Crustaceology. 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