Lycaea bovallii Chevreux 1900

Lycaea bovallii Chevreux, 1900

(Figs 1–3)

Lycaea Bovallii Chevreux, 1900: 157–159, pl. 18, fig. 3.— Stephensen 1925: 168–169, 230 (tab.).— Pirlot 1929: 136–137.— Belloc 1960: 21.

Lycaea bovallii. — Harbison & Madin 1976: 169, fig. 3A.— Thurston 1976: 388–389 (tab.), 434.— Stuck et al. 1980: 366.— Vinogradov 1990: 75, 94 (tab.).— Vinogradov 1991: 261 (tab.).— Vinogradov 1993: 45 (tab.).— Vinogradov 1999: 1195 (key).— Escobar-Briones et al. 2002: 367 (list).— LeCroy et al. 2009: 969 (tab.).

Lycaea bovalli. — Lin et al. 1996: 230 (tab.).— Vinogradov & Semenova 1996: 615.—Gasca 2004: 997, 998 (tabs.).— Gasca 2007: 118 (tab.).— Gasca 2009a: 89 (tab.).— Gasca 2009b: 66 (tab.).—Gasca et al. 2012: passim.— Gasca & Franco-Gordo 2014: 75 (list).

Lycaea bajensis Shoemaker, 1925: 46, figs 16–17.— Barnard 1930: 431–432, fig. 60.— Irie 1959: Table 4, 32 (tab.).— Harbison & Madin 1976: 169, fig. 5c.— Shih & Chen 1995: 171 (key), 174–176, fig. 113.— Lin et al. 1996: 230 (tab.).— Vinogradov & Semenova 1996: 615.— Zeidler 1998: 101.— Vinogradov 1999: 1195 (key).— Lowry 2000: 326 (list).— Escobar-Briones et al. 2002: 367 (list).— Gasca 2003a: 308 (tab.).— Gasca 2003b: 118 (tab.).— Gates et al. 2003: 321.—Gasca 2004: 997, 998 (tabs.).— Gasca 2007: 118 (tab.).— Gasca & Franco-Gordo 2008: 571.— Gasca 2009a: 89 (tab.), 91.— Gasca 2009b: 66 (tab.).— Gasca et al. 2009: 1497, 1501 (tabs.).— LeCroy et al. 2009: 969 (tab.).—Gasca et al. 2012: passim.— Gasca & Franco-Gordo 2014: 75 (list).— Hereu et al. 2020: passim. NEW SYNONYMY.

non [mis-identification].— Barnard 1931: 129–130.— Dakin & Colefax 1940: 124, fig. 214. (= Lycaea vincentii).

Lycaea gracilis Spandl, 1924: 30–32, fig. 6a-k.— Harbison & Madin 1976: 169–170, fig. 4c-d.— Vinogradov & Semenova 1996: 615. NEW SYNONYMY.

Lycaea Bovallioides Stephensen, 1925: 169, 230 (tab.), fig. 63.

Lycaea bovallioides. — Barnard 1930: 429 (key).— Evans 1961: 201.— Harbison & Madin 1976: 169, fig. 5A.— Thurston 1976: 388–309 (tabs.), 434.— Madin & Harbison 1977: 453 (tab.), 456.— Harbison et al. 1977: 470.— Laval 1980: 20 (tab.).— Stuck et al. 1980: 366.— Vinogradov 1990: 75–76, 94 (tab.).— Vinogradov 1991: 261 (tab.).— Vinogradov 1993: 45 (tab.).— Lin & Chen 1994: 118 (list).— Lin et al. 1995: 118, 122 (tab.).— Shih & Chen 1995: 171 (key), 176, figs 114– 115.— Lin et al. 1996: 230 (tab.).— Vinogradov & Semenova 1996: 615.— Vinogradov 1999: 1195 (key).— Lowry 2000: 326 (list).— Escobar-Briones et al. 2002: 367 (list).— Gasca 2003b: 118 (tab.).—Gasca 2004: 997, 998 (tabs.).— Vinogradov et al. 2004: 16, 25 (tab.).— Gasca 2007: 119 (tab.).— Gasca 2009a: 89 (tab.).— Gasca 2009b: 66 (tab.).— LeCroy et al. 2009: 969 (tab.).—Gasca et al. 2012: passim.— Nunes et al. 2013: passim.— Gasca & Franco-Gordo 2014: 75 (list).—Ambriz- Arreola et al. 2018: 59–61. NEW SYNONYMY.

Type material. Type material of Lycaea bovallii is in the Musée Océanographiqe de Monaco, Monaco (MOM). A holotype was not designated but a male from Hirondelle stn. 16 is illustrated by Chevreux (1900). Specimens were collected by the L’Hirondelle from the N.E. Atlantic, near the Azores [38°06’N 29°18’W], stn. 16, surface, 3 August 1885 (MOM 37 0019, 2 females, 3 males, in alcohol) and the N.E. Atlantic [48°19’N 19°30’W], stn. 30, surface, 27 August 1885 (MOM 37 0025, one juvenile, in alcohol) and [42°50’26”N 45°25’W], stn. 148, surface, 28 July 1887 (MOM 37 0025, one juvenile, in alcohol).

Type material of synonyms. The type material of Lycaea bajensis is in the USNM: the holotype male (USNM 52361) and 8 paratypes, 1 female, 7 males (USNM 52462). The type locality is the N.E. Pacific, Gulf of California. The holotype male (7 mm) and 8 paratypes are from off San Josef Island (“ Isla San José”). Shoemaker also lists additional material from this locality and from off Cape San Lucas (5) and from off Carmen Island (3) but most of these specimens could not be found in the USNM. However, there are two lots amongst the general collection, one male from Carmen Island (USNM 52463) and three poorly preserved specimens from Cape San Lucas (USNM 52464) that should be considered paratype material. Unfortunately, the latter lot has only one specimen, a male that could be referred to L. bajensis; the remaining two specimens are females that are more readily identified with L. pulex and L. vincentii. An examination of this material has confirmed the above synonymy.

Several syntypes of Lycaea gracilis are in the Naturhistorisches Museum Wien, in alcohol (NHMW 20163). Several specimens were collected from the Red Sea; Pola stns. 22–25, 64, 82, 108, 118, 136, and a male is illustrated by Spandl (1924). The long dactylus of P4, the relatively long peduncle of U1, the morphology of G1 and G2, and of the male A1 are all characteristic of L. bovallii.

Several syntypes of Lycaea bovallioides (4–5 mm) are in the NHMD, all labelled “Type”, in alcohol. Specimens were collected by the Thor from surface waters in the Mediterranean Sea, stn. 163 [37°52’N 26°22’E], 3 August 1910 (1 female, NHMD-83719, previously CRU-5875) and stn. 718 [36°13’N 13°53’E], 17 May 1913 (1 female, NHMD-86792, previously CRU-9293), and from the N.E. Atlantic, near Madeira, stn. 398 [36°48’N 14°22’W], 26 October 1911 (1 female, 1 male, NHMD-86789, previously CRU-9290); stn. 399 [34°23’N 15°31’W], 26 October 1911 (3 females, 3 males, NHMD-86790, previously CRU-9291) and stn. 400 [32°10’N 17°20’W], 30 October 1911 (12 males, NHMD-86791, previously CRU-9292). The figured male is from stn. 400 (Stephensen 1925) and is here designated the lectotype, the remaining syntypes become paralectotypes. An examination of this material has confirmed the above synonymy.

Material examined. Type material of Lycaea bajensis and L. bovallioides as detailed above and the following. In NHMD: N. Atlantic, 9 females, 5 males (6 lots), Dana stns 1142 vii, 1145 vii (228137–8), 1231 v (228141), 4000 v (228231), and Thor stns 385, 399. S.W. Atlantic, 3 females, 1 male, Dana stn. 3997 v (228228). Indian Ocean, from Sumatra to South Africa, 18 females, 7 males (17 lots, all Dana), stns 3821 v, 3843 i, 3844 iii-iv, 3851 ii-iv (228175–9, 228181, 619434), 3854 i (228183), 3918 v (228189), 3920 viii, 3921 ii, 3921 vii (228193–5), 3928 iii (228203), 3931 iii (619436), 3937 ii (228216), 3959 iv (228223), 3843 iv (228255). Tropical Pacific, 20 females, 16 males (20 lots, all Dana), stns 3548 v, 3553 i (228143–4); 3561 x (228146); 3563 ii, 3569 I, 3569 iv, (228148–50); 3579 v, 3584 ii (228152–3); 3584 vi, 3584 vii (228155–6); 3585 v (228158); 3587 ix (228161); 3588 ii, 3588 iii, 3602 v, 3611 iv 3611 vi, 3626 iv, 3663 ix–x (228163–70). Tasman Sea, 1 female, Dana stn. 3665 iv (619246). Mediterranean Sea, 19 females, 11 males (11 lots), Dana stns 4050 xxi-iii, 4050 v (228233–6), and Thor stns 10, 143, 144, 163, 182, 183, 297. In SAM and SAMA (part): Meiring Naude collections from S.W. Indian Ocean, off South Africa, between Kosi Bay and just south of East London, 97 females, 63 males (53 lots), mostly 200–0 m, few 528–0 m. In SAM: S.W. Atlantic, off South Africa, 19 females, 8 males (7 lots). In SAMA: N.E. Indian Ocean, off northern Western Australia, Ningaloo Reef and Dampier Archipelago region, 8 females, 19 males (9 lots), C12558–66. Tasman Sea, north of Sydney, 1 female, 1 male (2 lots), C5264–5. Japan, Ryukyu Islands, 2 females, 1 male (2 lots), C12567–8. S.W. Atlantic, off Brazil [31°05’S 49°50’W], 2 males, C12569. In USNM: N.W. Atlantic from French Guiana in the south, north to Georges Bank, off Massachusetts, 38 females, 17 males (28 lots), 106183, 181803, 1154670, 1154682, 1171017, 1178027, 1198726, 1241233, 1241235, 1241238, 1241240, 1241242, 1241245, 1241286, 1242779, 1242784–5, 1242787, 1242793, 1242807, 1242810–11, 1246891, 1246972–3, 1246977–8, 1247116, 1247126. S.W. Atlantic, off Brazil [08°S 30°W], 3 females, 2 males (2 lots), 1246990, 1247117. N.E. tropical Pacific, off Costa Rica and Nicaragua, 5 females, 20 males (3 lots), 1242773, 1242776, 1242778. N.W. Pacific, Japan, Philippines and China Sea region, 12 females, 26 males (10 lots), 1242774–5, 1242777, 1242796, 1242798–800, 1242803–4, 1246961. N.W. Indian Ocean, near Mauritius [19°34’N 62°14’09”E], 1 female, 1246891.

Diagnosis. Body length up to about 5.0 mm for females and 7.0 mm for males, but ovigerous females of about 3.0 mm have been noted. Head of females relatively large, much deeper than long (about 1.5 x), as long as first 5 pereonites combined, sometimes slightly shorter or longer. Head of males more rounded, slightly deeper than long (about 1.3 x), almost as long as first 5 pereonites combined. Buccal mass barely protruded below head. Callynophore of A1 of males with acute antero-distal corner extending at right angles well above following article; postero-distal corner small, rounded, separated from following article by distinct notch. G1 and G2 sub-chelate, morphologically similar, G2 slightly longer than G1; basis of G1 slightly broader and shorter than G2; carpus rectangular with sharp postero-distal tooth, reaching just past base of dactylus; propodus with postero-distal corner produced posteriorly to dactylus; carpus and propodus with small serrations on distal and posterior margins; dactylus slender, length 0.5–0.6 (females), 0.6–0.8 (males) x propodus. P3–6 with relatively long, slender dactylus, those of P3 and P4 at least 0.5 x as long as propodus or longer. P3 and P4 morphologically similar, P4 slightly longer than P3; merus marginally inflated anteriorly, slightly longer than propodus, about 0.6 x basis; carpus length about 0.6–0.7 x propodus. P5 length about 1.2 x P 4 in females, 1.4 x P 4 in males and about 1.4 x P 6 in both sexes; basis rectangular, length about 2.4 x maximum width; merus marginally inflated anteriorly, slightly longer than propodus, about 0.6–0.7 x basis; carpus length about 0.7 x propodus. P6 basis oval-shaped, length about 2 x maximum width, slightly shorter than basis of P5; merus relatively narrow; merus, carpus and propodus similar in relative lengths to P5; anterior margin of carpus and propodus, and antero-distal corner of merus, slightly serrated. P7 basis with bulging posterior margin, wider in males with length about 1.5 x maximum width, in females length almost 2 x width, about 0.6–0.7 x basis of P6; remaining articles together very short, less than 0.2 x basis; propodus with small, rounded, antero-distal corner; dactylus sharp, hook-like. U1 peduncle relatively long, more than 3.0 x length of exopod; rami relatively slender, equal in length. U2 endopod usually fused with peduncle. Telson length about equal to width at base in males, slightly longer in females.

Remarks. The validity of Lycaea bovallii has been in doubt in the past and Vinogradov et al. (1982, 1996) consider it a synonym of L. pulex, along with L. bovallioides and L. bajensis. However, like Harbison & Madin (1976), it is here regarded a valid species, distinguished from L. pulex by the relatively longer peduncle of U1, in that the endopod of U2 is usually fused with the peduncle, and by the relatively longer dactylus of P4 (and other pereopods). Males are further distinguished by the morphology of the callynophore of A1 as detailed above. Lycaea gracilis, L. bovallioides, and L. bajensis, cannot be clearly distinguished from L. bovallii, as demonstrated above and are thus considered junior synonyms. However, this species is very similar to L. vincentii in the relatively long peduncle of U1 and in the morphology of the callynophore of A1 of males. Regarding the latter character, the main difference is in the position of the antero-distal bulge as detailed above. Apart from the relatively longer dactylus of P4 (and other pereopods), it is most readily distinguished from L. vincentii by the fused endopod of U2, although this is not always easy to determine. Also, the buccal mass is not protruded as much below the head and the remaining articles of P7, distal to the basis, are together relatively short, less than 0.2 x the basis length compared to 0.3 x, or more, for L. vincentii, although this character may be variable.

In view of the above synonymy, especially that of L. bajensis, Lycaea bovallii has become one of the more common species of Lycaea in the N.W. Atlantic Ocean.

Lycaea bovallii has been recorded with the following salps (as L. bovallioides), Cyclosalpa pinnata (Forsskål, 1775), Pegea socia (Bosc, 1802), P. confoederata (Forsskål, 1775), Iasis cylindrica (Cuvier, 1804) and S. maxima Forsskål, 1775 (Madin & Harbison 1977). It has also been recorded with the pteropod Corolla spectabilis Dall, 1871 (Harbison et al. 1977).

Distribution. Determining the distribution of this species is problematical, mainly because of its past confusion with L. pulex. Material in the USNM is mainly from the N.W. Atlantic Ocean, as might be expected. Considering the above synonymy, more reliable records are as follows. In the Atlantic Ocean: from about 40°N, throughout the tropical regions, and as far south as 31°S, off Brazil. In the Pacific Ocean: mainly from the tropics off Chile and Peru, the Gulf of California, the China Sea and near Japan, and the Tasman Sea, off eastern Australia. In the Indian Ocean: mainly from the tropical south-west and also the Red Sea. The SAMA also has several specimens collected from off the north-western coast of Western Australia. It has also been recorded from the Mediterranean Sea. Most of these records are from near the surface.