Trimma benjamini Winterbottom, 1996, Winterbottom, 1996

Winterbottom, Richard & Hoese, Douglass F., 2015, A revision of the Australian species of Trimma (Actinopterygii, Gobiidae), with descriptions of six new species and redescriptions of twenty-three valid species, Zootaxa 3934 (1), pp. 1-102: 14-19

publication ID

http://dx.doi.org/10.11646/zootaxa.3934.1.1

publication LSID

lsid:zoobank.org:pub:11C2A2CB-30B3-4694-B379-AE9D47332F0C

persistent identifier

http://treatment.plazi.org/id/5519879A-B80E-F357-FF1F-FB7E6A56E988

treatment provided by

Plazi

scientific name

Trimma benjamini Winterbottom, 1996
status

 

Trimma benjamini Winterbottom, 1996  

Ringeye Pygmygoby

Figs. 5 View FIGURE 5 , 6 View FIGURE 6 , Pl. 1 C)

Trimma benjamini Winterbottom, 1996: 57   , figs. 1–5 (Siquijor Id, Philippines); Allen et al., 2004: 329; Hayashi & Shiratori, 2003: 41 (x 2); Yano, 1998: 27; Kimura & Matsuura, 2003: 191; Kuiter & Tonozuka, 2004: 703; Myers, 1999: 163; Randall, 2005: 552; Randall et al., 1997: 409 (Great Barrier Reef); Suzuki et al., 2007 b (first record from Japan); Allen & Erdmann, 2012: 935 (Borneo to Marshalls, Tonga).

Trimma   sp.: Burgess et al., 1990: 577.

Trimma   sp. A: Kuiter, 1992: 237 A.

Trimma   sp. 8: Senou et al., 2004: 112 (x 2).

Trimma   sp. 5 (DFH 4): Allen, 1993: 88 ( Ashmore Reef).

Trimma   sp. 2 (DFH 4): Randall et al., 1997: 409 (Great Barrier Reef).

Australian Material. Queensland. Cape Melville, reef off: AMS I. 20774 -112, 29(12–20), 0– 8 m. Escape Reef: AMS I. 22576 -035, 12 (19–23), 34 m; AMS I. 22578 -061, 19 (17–24), 6–10 m; AMS I. 22578 -063, 2 (12–13), 6–10 m; AMS I. 22580 -018, 31 (13–21), 37 m; AMS I. 22581 -045, 27 (11–23), 10–14 m; AMS I. 22583 -041, 4 (16–21), 29 m; AMS I. 22587 -026, 23 (13–22), 40 m; AMS I.22611, 3(19–20), AMS I.22612, 3(11–21), 20 m; AMS I. 22613 - 0 44, 45(13–22), 27 m; AMS I. 22619 -040, 6 (12–22), 19–22 m; AMS I. 22620 -017, 2 (11–21), 38–50 m; AMS I. 22621 -039, 8 (14–20), 5–8 m; AMS I. 22627 -015, 4 (18–21), 56–60 m; AMS I. 22637 -048, 23 (10–24), 3– 15 m. Lizard Id: AMS I. 19607 - 110, (20), 5–6 m; AMS I. 20730 -013, 4 (18–24), 20–22 m; AMS I. 21422 -093, (20), 1– 8 m. McGillivray Reef: AMS I. 18805 -006, 2 (16–24), 11–15 m; AMS I. 19482 -141, 11(9–22), 3– 25 m. North Direction Id: AMS I. 19442 -077, (21), 12 m. One Tree Id: AMS I. 15637 -045, (23), 23–26 m; AMS I. 15682 -060, 3 (17–21), 30 m; AMS I. 15684 -057, 2 (18–24), 30 m; AMS I. 17929 -017, 4 (22–25), 33 m; AMS I. 20206 -015, 14 (11–24), 22– 30 m. Pikie Reef: WAM P. 25322 -008, (22). Raine Id: AMS I. 20757 -080, 12 (17–20), 0–20 m; AMS I. 20775 -119, 40(9–22), 0– 20 m. Yonge Reef: AMS I. 19445 -097, 26 (15–23), 15 m; AMS I. 19454 -064, 28 (13–23), 1–18 m; AMS I. 19454 -091, 3 (18–22), 1–18 m; AMS I. 19456 -073, 14 (14–22), 5–15 m; AMS I. 19472 -094, 4 (18–23), 7–15 m; AMS I. 19481 -043, 8 (12–23), 2–6 m; AMS I. 20784 -023, (22), 1–15 m; ROM 39332 View Materials , 2(16–20), 8– 13 m. Tijou Reef: AMS I. 20779-147, 100 (10–26), 0–25 m; AMS I. 20956 -018, 13 (14–27), 3– 12 m. Townsville, reef off: AMS I. 20965 -012, 52 (15–22), 10– 20 m. Western Australia: Ashmore Reef: NTM S. 12318 -059, (21), 14– 16 m. Rowley Shoals: WAM P. 28026 -025, (22), 33–40m; WAM P. 28034 -016, (17), 33– 40 m.

Other material. American Samoa: AMS I. 21986 -002, 6 (12–25), 15 m; AMS I. 21987 -002, 6 (12–25); AMS I. 21988 -001, 2 (17–19), 58–64 m; BMNH, 2 (21 – 21); BMNH [RL 8 – 3], (21), 30–40 m; BPBM (uncatalogued), 2 (20–21), 37 m. Micronesia: Ulithi Atoll, BPBM [uncatalogued], (18), 10 m. Fiji: Astrolabe Reef: ROM 40108 View Materials , 10(11–17), 16 m; ROM 46104 View Materials , 8(13–25); ROM 46105 View Materials , 6(9–17), 9–11m; ROM 46106 View Materials , 19(11–18), 23–30 m; ROM 46109 View Materials , (18); ROM 46110 View Materials , 7(9–20), 3–10 m; ROM 46111 View Materials , 14(10–21), 5–8 m; ROM 46112 View Materials , (23); ROM 46113 View Materials , 29(11–22); ROM 46114 View Materials , 31(10–21), 5–10 m; ROM 46115 View Materials , 20(14–19), 2–7 m; ROM 46116 View Materials , 45(10–20), 27 m; ROM 46117 View Materials , 7(15–18), 25–30 m; ROM 46118 View Materials , 10(9–22), 30 m; USNM 313325, 18(13–20), 0–40 m; USNM 313330, 3(11–14), 0– 40 m. Bagilai Is: ROM 40058 View Materials , 2(18–24). Bega: AMS I. 18419 –016, 3 (18–20), 27– 37 m. Dawson Reef, USNM 236716, 34(16–22), 0– 14 m. Makuluja Reef, ROM 46119 View Materials , 20(10–18), 15– 20 m. Malamala Id, USNM 236725, 6(21–26), 0– 8 m. Malolo Reef, USNM 236721, 7(14–40), 0–18 m; USNM 236731, 17(15–26), 21–29 m; USNM 256503, (16), 21– 29 m. Matuku Id, USNM 236753, 2(18–23), 23 m; USNM 243043, 10(17–22), 20–23 m; USNM 243199, 14(17–20), 0–12 m; USNM 256497, (19), 0– 12 m. Naviti Id, USNM 236770, 2(13–18), 24– 30 m. Ono-Ilau Id: ROM 1323 CS, 5 (17–20), 43 m. Rotuma, USNM 283030, 12(11–22), 0–41 m; USNM 285695, 3(17–23), 0– 41 m. Suva: AMS I. 18352 -002, 2 (15–18); AMS I. 18352 -033, 3 (19–20); AMS I. 18353 -034, (26), 6 m; AMS I. 18354 -105, 18(10–25), 6 m; AMS I. 18356 -009, (23); ROM 46120 View Materials , 8(15–20), 10–15 m; ROM 46121 View Materials , 58(7–21), 10–14 m; ROM 46122 View Materials , 2(18–20), 10– 14 m. Totoya Id, USNM 236764, 5(17–27), 30– 34 m. Viti Levu, USNM 243092, 40(10–22), 9–37 m; USNM 243193, 3(14–20), 9– 12 m. Viwa Id, USNM 241782, (14), 21– 30 m. Indonesia: Ambon, Moluccas Ids: WAM P. 25236 -013, (16), 5 m. Raja Ampat: ROM 84932 View Materials , 2(10 – 5), 2–5 m; ROM 85065 View Materials , 5(14–21), 50 m; ROM 85077 View Materials , 18(10–20), 15–50 m; ROM 85107 View Materials , 4(15–18), 18–23m; ROM 85124 View Materials , 6(9–18), 15–31 m; ROM 85154 View Materials , 2(13 – 13), 20–23 m; ROM 85227 View Materials , 1(22), 12–16 m; ROM 85252 View Materials , 16(10–21), 12–26 m; ROM 85277 View Materials , 2(9–12), 15–22 m; ROM 85287 View Materials , 8(9–19), 2–28 m; ROM 85300 View Materials , 1(17), 9–12 m; ROM 85378 View Materials , 5(16–20), 14–18 m; ROM 87453 View Materials , 2(17 – 17), 15– 18 m. New Caledonia: ROM 63934 View Materials , 18(18–28), 18–26 m; ROM 63935 View Materials , 19(19–25), 18.3–27.4 m; ROM 64328 View Materials , 10(15–29), 6– 11 m. Palau: Main Ids: BPBM [uncatalogued], 4 (13–16), 33–52 m; BPBM [uncatalogued], 4 (13–21); ROM 74942 View Materials , 84(11–22), 12–21 m; ROM 80370 View Materials , 54(12–20), 14–18.4 m. Helen Reef: ROM 83221 View Materials , 39(9–20), 15– 28 m. Papua New Guinea: Kavieng (New Ireland): WAM P. 28186 -010, 6 (13–21). Kiriwinna (Trobriand Ids): AMS I. 17102 -074, (18), 0– 4 m. Manus Id: WAM P. 27824 -042, 2 (16–17), 19–20 m; WAM P. 27825 -061, 5 (13–20), 10–40 m; WAM P. 27826 -086, 21 (9–19), 35– 41 m. Rabaul: ROM 88132 View Materials , (15), 12.5 m; WAM P. 28172 -042, 4 (16–27); WAM P. 28174 -025, 10 (17–25), 25– 30 m. Rapa Point: WAM P. 28165 -012, 4 (14–19), 32– 40 m. Philippines: Anilao: AMS I. 21907 -007, (22), 21 m. Caban Id: AMS I. 21914 -020, 25 (13–22), 9–25 m; AMS I. 21918 -030, 50 (13–27), 11–29 m; AMS I. 21922 -020, 3 (17–19), 18– 21 m. Cebu: ROM 1309 CS, 5 (19–24), 3– 11 m. Negros Id: ROM 53034 View Materials , 36(9–23), 3–5 m; ROM 53035 View Materials , 26(9–22), 6– 12 m. Puerto Galera (Mindoro): LACM 42491 - 70, (20). Siquijor Id: ROM 53028 View Materials , 11(18–23), 5– 12 m. Sombrero Id: AMS I. 21908 -013, 31 (12–22), 1– 34 m. Solomon Ids: Florida Id: AMS I. 17500 -029, 3 (16–19), 20–30 m; AMS I. 20395 -009, (16). Guadalcanal: AMS I. 17524 -006, (22), 35 m; AMS I. 27020 -004, (17), 32 m; ROM 0840CS, 14 (11–29), 8 m; ROM 46049 View Materials , 15(10–20), 18– 22 m. Santa Cruz Id: AMS I. 39031 -069, 5 (15–22), 28 m; AMS I. 39034 -016, 7 (15–27); AMS I. 39034 -016, 7 (15–27), 5 m; AMS I. 39036 -055, 6 (12–21); AMS I. 39036 -055, 6 (12–21), 35 m. Savo Id: AMS I. 17489 -016, 6 (16–21), 20 m; AMS I. 17491 –040, 6 (17–21), 10 m. Tonga: Vava'u: ROM 60683 View Materials , 2(19–21), 36–40 m; ROM 60683 View Materials , 2(19–21), 37–40 m; ROM 60684 View Materials , 4(17–22), 25 m; USNM 295226, 10(18–22), 0– 16 m. Vanuatu: Bogacio Id: AMS I. 17477 -031, 6 (17–19), 55 m. Efate Id: AMS I. 17472 -068, 3 (16–18), 10 m; AMS I. 18434 -006, (21). Off Emae: AMS I. 17475 -054, 9 (17–24), 25 m. Futuna Id: AMS I. 20792 -039, (18). Namuka Id: AMS I. 37331 -022, 2 (17–20), 35 m. Vietnam: Nha Trang: ROM 73186 View Materials , 2(17), 23–29 m; ROM 73187 View Materials , 2(15–16), 21– 30 m.

Diagnosis. A species of Trimma   with a deep concave, ‘U’-shaped interorbital trench; bony interorbital about half pupil-diameter in width; a slight groove posterodorsal to the eye; predorsal midline naked, sides of nape covered with small scales to or near to the eyes; cheek and operculum naked; pectoral-fin base with oval, moderatesized scales; prepelvic area with 5–6 rows of cycloid scales, isthmus naked anteriorly, one slightly enlarged scale on the membrane between the bases of the pelvic fins; central 6–13 pectoral rays branched, uppermost and lowermost rays unbranched; fifth pelvic-fin ray branched once dichotomously and about 60–75 % length of the fourth; other rays usually with one sequential branch, basal membrane connecting the two pelvic fins only at the base; second dorsal spine longest, usually filamentous, reaching at most to midway along the base of the second dorsal fin in Australian material; second dorsal fin-rays usually I 10; anal-fin rays usually I 9; nape crest fleshy from the first dorsal origin to above the operculum; head and body red in life, yellow in preserved material, head with a thin vertical grey (bluish grey in life) bar from the anteroventral margin of the eye to the middle of the jaws and a second bar extending more-or-less vertically downward onto cheek from about the middle of the eye (often persisting in preserved material as faint grey bars); orbit rimmed with a thin band of melanophores, more prominent dorsally (faded in preserved material).

Description. The description is primarily based on specimens from the Great Barrier. Dorsal fin VI + I 10 (I 9 in 8.5 % of specimens, rarely I 8 or I 11 in <1 % of specimens, mean = I 9.9, n = 358), second spine longest, reaching to at least to second segmented dorsal ray to just beyond middle of second dorsal fin when adpressed, first ray of dorsal fin branched or unbranched (half of 30 specimens between 15 and 24 mm SL examined had a branched ray), anterior element of last element branched; anal fin I 9 (I 8 or I 10, in 7.3 % of specimens and rarely I 7 in <1 % of specimens, mean = I 8.9, n = 358), first ray usually unbranched, anterior element of last ray branched; pectoral-fin rays 17–20, usually 18–19 (in 93 % of specimens examined), but varying a little geographically (see Table 1), reaching just posteriorly to a vertical in line with anal spine or first 2–3 segmented anal rays, pectoral fin with 6–13 central rays branched, upper 3–5 rays usually unbranched and lower 4–8 usually unbranched, branching increasing with size (6–10 rays branched in specimens less than 20 mm SL to 8–13 rays branched in larger specimens); pelvic fin I 5, rays usually with 1 sequential branch, sometimes second, third or fourth segmented ray branched twice (with 3 terminal tips), fifth ray with a single dichotomous branch (two terminal tips), 65–75 % the length of the fourth ray, which reaches posteriorly to anterior few elements (second to fourth segmented ray) of anal fin; fraenum absent, basal membrane variably present in preserved specimens from just connected to basal 10–20 % of fifth ray in well preserved material, but often torn. Lateral scales 21–25, usually 23 or 24 (in 79 % of individuals), rarely 21 (in 2 % of individuals), mean = 23.5, n = 105 for Australia and 23.2, n = 102 for other localities; transverse scale count usually 7–8, (rarely 9 in <10 % of individuals), mean = 7.6, n = 105 for Australia and 7.8, n = 88 for other localities; scales on pectoral-fin base, breast, and anteriormost scales on midline of belly cycloid, remaining scales ctenoid; midline of nape naked, sides with scales extending anteriorly to just behind eye; scales below posterior end of second dorsal fin with small accessory scales; pectoral-fin base usually with 4 vertical rows of cycloid scales, 1 horizontal row dorsally and 3 ventrally, none of scales prominently enlarged or elongate; prepelvic area with 5–6 rows of cycloid scales, a single enlarged median scale and sometimes one or two smaller median scales covering membrane between pelvic fins. Teeth in lower jaw consist of an enlarged outer row of curved, spaced canines and an inner row of similar, but smaller canines (about two thirds size of outer teeth), with irregular rows of small conical teeth in between; outer row of teeth in the upper jaw similar to those of the lower jaw, with small irregular inner rows of teeth. Tongue rounded to truncate, about two-thirds pupil diameter in width, with pointed tip ventrally. Gill opening extending anteroventrally to below or just behind mid-pupil; gill rakers on first arch 3–5 + 13–15 (mean = 17.7 based on 58 specimens; Winterbottom (1996) reported 3–4 + 13–14 (mean = 3.8 + 13.8 )). Anterior nares at end of short tube just above upper lip; posterior nares pore-like with a raised rim, 1 nares diameter from anterior and 1–2 nares diameters from fleshy rim of eye. Nasal sac slightly elevated. Interorbital trough moderate and ‘U’-shaped, posterodorsal trough represented by a shallow groove between the epaxialis muscle along posterodorsal margin of eye and bony rim of orbit; bony interorbital width about half to one third pupil diameter; epaxialis musculature reaching to a point in line with a vertical with posterior margin of the pupil. Posterior abdominal/anterior caudal vertebral pattern is Type B.

Colour pattern. Freshly collected. (From slides of a 22 mm SL specimen from the Great Barrier Reef, Fig. 5 View FIGURE 5 ). Snout and lips red, rest of head and body normally red to reddish orange, rarely yellowish orange; a thin blue bar along posteroventral margin of eye expanding into a thin bar extending ventrally to behind jaws; anteriorly a thin blue line from anteroventral margin of eye extending to just behind middle of lips; often with a third blue mark just behind posterior nostril. Pupil surrounded by a thin yellow to gold circle; iris reddish orange, often golden ventrally. Body often with melanophores arranged in vertical wavy lines along centers of scales; caudal peduncle with faint grey stripes formed by dense subcutaneous dark pigment. Dorsal fins largely clear, with a thin yellow to red stripe just above base of fin; a second thin stripe just below middle of fin, most prominent on second dorsal fin. Second dorsal fin yellowish to pale orange above stripes, with a thin white to bluish distal margin. Anal fin red to reddish orange with thin blue to white distal margin. Caudal fin reddish orange to yellow basally grading posteriorly to a translucent grey, distal margin often pale blue. Pectoral and pelvic fins white to pale grey. Live specimens from type locality ( Philippines), and those from Great Barrier Reef (Pl. 1 C) overall bright red with underlying dusting of brown chromatophores. Specimens from Palau and some other parts of western Pacific may be much yellower (although adult males exhibit a bright red head).

Preserved. Head and body largely plain straw coloured; bar below eye often persisting as a grey band; fins largely pale, except dorsal fins often with scattered melanophores.

Variation. Branching of pectoral rays increases significantly with size (p = 0.01, based on 70 specimens selected at random from 15–25.5 mm SL). Specimens below 20 mm SL usually have 5–7 branched rays, while specimens above 20 mm SL usually have 8–11 branched rays. The ranges of counts are, however, highly variable at all sizes.

There is considerable variation in the length of the second filamentous dorsal spine. In most Australian material the spine is typically short, rarely reaching beyond the middle of the second dorsal fin when adpressed. In specimens from the Philippines, the spine often reaches beyond the end of the second dorsal fin.

Pectoral ray counts show significant variation between samples ( Table 1). Samples from Fiji average higher pectoral ray counts (19.0) than other localities. Analysis of Variance on samples of> 20 specimens indicate that the samples are not from a single population (p <0.05). Samples from the Great Barrier Reef show little variation, with means between 18.4 and 18.6.

The blue lines below the eye are absent in photos of some fresh specimens from some localities (Yonge Reef, Queensland, Solomon Islands and Samoa). Other photos from these localities show the bar below the eye. The lines are faint in photos of a fresh specimen from New Caledonia. In others, the lines are sometimes incomplete, particularly ventrally on the cheek. The overall body colouration varies from bright red to a red head and reddish orange body. The red colouration fades rapidly in preserved material.

Sex ratios were normally about even, with a total of 155 males and 193 females examined; however, because of the slightly larger size one would expect a bias of more males. The maximum size known is 29 mm SL from New Caledonia (with a 28 mm SL specimen from Lizard Island), but for most samples the maximum size is around 20–25 mm SL. Males average about 1 mm SL larger in most samples (p <0.05) and reach a size larger than females of 2–3 mm SL in any sample. In some small individuals the urogenital papilla of females is pigmented like that of males and it is possible that some individuals change sex.

Etymology. Named for Peter Benjamin, Toronto entrepreneur, whose company, Benjamin Films Ltd., provided RW with decades of free film and processing, and who participated in two ROM collecting expeditions.

Distribution. Trimma benjamini   is abundant on the Great Barrier Reef, currently known from One Tree Island on the southern part of the reef and from Pixie Reef off Cairns to Ashmore Reef on the northern part of the reef. It is also found on islands of the Coral Sea and Ashmore Reef in the Timor Sea. It is normally confined to offshore reefs along the eastern edge of the Great Barrier Reef and only rarely found on mid reefs near Lizard Island and never from coastal reefs. It is also known from Rowley Shoals and Ashmore Reef off Western Australia. In Australia it is most abundant in depths of 10–30 metres, but is found from 1– 37 m. Outside of Australia the species was found in similar clear water reefs to depths of 68 m. The species is known from Vietnam and Western Australia eastward to the Marshall Islands and Samoa ( Fig 6 View FIGURE 6 ). Specific locations outside of Australia include Palau, Moluccas, Papua New Guinea, Marshall Islands, Vietnam, Fiji, Philippines, New Caledonia, Solomon Islands, Tonga, Vanuatu and Samoa.

Comparisons. Among the species of Trimma   which lack scales in the predorsal midline and have a uniform body colouration (i.e. lacking light spots, saddles, streaks or distinct stripes, at least when preserved), only two other species have an interorbital trough as deep as it is wide. Those species, Trimma corallinum ( Smith 1959)   and T. omanensis Winterbottom, 2000   , are currently known only from the western Indian Ocean, have 20–21 (vs. usually 18–19) pectoral rays, a full basal (vs. one-tenth) membrane, a fifth pelvic-fin ray sub-equal (vs. 60–75 %) to the fourth, and a well-developed trough (vs. a slight groove) posterodorsal to the eye. Pale specimens without a bar under the eye can be similar in colouration to T. preclarum   (which has faint stripes on the caudal peduncle). That species differs in having predorsal scales crossing the midline and lower dorsal ray counts.

The analysis of the CO 1 gene suggests there are at least two haplogroups, one (n = 4) from the main islands of Palau, and the other from a more southerly part of the Pacific (three from the Great Barrier Reef, one from New Caledonia, and two from Helen Reef in the South West Islands of Palau) separated by approximately 5.6 % of the CO 1 genome ( Winterbottom et al., 2014). The Helen Reef specimens are separated by about 0.3 % from those from Australia and New Caledonia. No specimens were available for genetic analysis from the type locality (Siquijor Island, Philippines).

Discussion. Winterbottom et al. (2011) found that the South West Islands ( Palau) population identified as this species had maximum age of 140 days with an average pelagic larval duration of 33.9 ± 4.3 days, or 24.2 % of the maximum lifespan. Estimates of daily mortality rate ranged between 2.9 % – 6.3 %.

This species has been informally referred to as Trimma   RW sp. 18 or T. DFH sp. 4 prior to its formal description.

TABLE 1. Pectoral ray counts of select populations of Trimma benjamini.

  17 18 19 20 21    
  2 33 47     85  
    24     36  
  1 12 10     23  
  - 15 13     29  
          13  
    13     19  
  1 14       18  
    18     26  
  3 12       24  
  5 24 27     56  
WAM

Western Australian Museum

ROM

Royal Ontario Museum

NTM

Northern Territory Museum of Arts and Sciences

BPBM

Bishop Museum

USNM

Smithsonian Institution, National Museum of Natural History

LACM

Natural History Museum of Los Angeles County

Kingdom

Animalia

Phylum

Chordata

Class

Actinopterygii

Order

Perciformes

Family

Gobiidae

Genus

Trimma

Loc

Trimma benjamini Winterbottom, 1996

Winterbottom, Richard & Hoese, Douglass F. 2015
2015
Loc

Trimma

Senou 2004: 112
2004
Loc

Trimma

Randall 1997: 409
1997
Loc

Trimma benjamini

Allen 2012: 935
Randall 2005: 552
Allen 2004: 329
Kuiter 2004: 703
Hayashi 2003: 41
Kimura 2003: 191
Myers 1999: 163
Yano 1998: 27
Randall 1997: 409
Winterbottom 1996: 57
1996
Loc

Trimma

Allen 1993: 88
1993
Loc

Trimma

Kuiter 1992: 237
1992
Loc

Trimma

Burgess 1990: 577
1990