Littorininae, Children, 1834

Reid, David G., 2011, 2974, Zootaxa 2974, pp. 1-65 : 9-10

Key to the Littorininae of the tropical and warm-temperate eastern Atlantic

There are no known morphological synapomorphies by which to diagnose the genus Echinolittorina . The shells are convergent with those of several other littorinid genera. Therefore this key includes all 14 species of the subfamily Littorininae that have been recorded from the tropical and warm-temperate eastern Atlantic Ocean (including the Mediterranean Sea), belonging to the genera Echinolittorina , Afrolittorina , Tectarius , Littoraria , Littorina and Melarhaphe . Discrimination is based primarily on characters of the shell, with the addition of diagnostic characters of the penis and pallial oviduct when these are easily observable. References to figures of shells, penes and oviducts of all species are given, together with information on distribution and habitat. All species occur only on rocks in the littoral fringe unless otherwise noted.

1 Nodulose or granulose, over entire shell or early whorls only................................................... 2

- Spiral ribs or grooves, or smooth......................................................................... 5

2 1 pale band in aperture; 0 or 3 rows of small nodules on last whorl; no threads between spiral ribs; penis without glands; pallial oviduct with albumen and capsule glands in two successive spirals; Azores, Madeira, Canary Is, Cape Verde Is..................................nodulose form of Tectarius (Liralittorina) striatus ( King & Broderip, 1832) ( Reid 1996: figs 8–10)

- 1–2 pale bands in aperture; 0 or 4–7 rows of nodules on last whorl; usually 1–3 threads between spiral ribs; penis with mamilliform gland and glandular disc; pallial oviduct with albumen and capsule glands in same spiral; tropical West Africa, St Helena, Ascension I................................................................................... ……3

3 Nodules usually large, axially aligned, black when eroded; tropical West Africa, islands in Gulf of Guinea; sometimes in estuaries............................................................................ E. granosa ( Figs 16, 17)

- Nodules small or absent, not conspicuously axially aligned, paler than background; St Helena, Ascension I............... 4

4 Last whorl with 4–7 rows of small pointed nodules of equal size; Ascension I.................... E. miliaris ( Figs 20, 21)

- Last whorl with 0 or 4 rows of small rounded nodules, anterior 2 rows larger; St Helena.............................................................................................. ……nodulose form of E. helenae ( Figs 22, 23)

5 Small, <9 mm; smooth or with single incised line at periphery; penis without glands................................ 6

- Up to 33 mm; spiral ribs or grooves or smooth; penis with or without glands (but, if shell smooth, then penis with glands).. 7

6 Smooth with single incised line at periphery; brown with white spots or lines; narrow, deep pseudumbilicus. Pallial oviduct with albumen and capsule glands in 1 spiral. Tropical West Africa. Rocks and algae............. E. peregrinator ( Figs 2, 3)

- Entirely smooth; grey to brown, sometimes dark dorsal zone and paler base, no white spots; no pseudumbilicus. Pallial oviduct with glands in 2 successive spirals. NW Europe to Mauretania, Azores, Madeira, Canary Is........................................................................... Melarhaphe neritoides ( Linnaeus, 1758) ( Reid 1989: figs 1l, 6l, 9l)

7 Last whorl with 15–20 sharp, equal ribs without intervening threads; grey with fine brown speckling; penis without glands; pallial oviduct with albumen and capsule glands in 2 successive spirals; Azores, Madeira, Canary Is, Cape Verde Is................................. striate form of Tectarius (Liralittorina) striatus ( King & Broderip, 1832) ( Reid 1996: figs 8–10)

- Penis with glands; pallial oviduct with albumen and capsule glands in 1 spiral..................................... 8

8 No pale band in aperture; penis with glandular disc only.............................................. Littoraria 9

- Penis with1 or more mamilliform glands and sometimes glandular disc.......................................... 10

9 To 17 mm; 22–33 grooves on last whorl; cream with dense spiral lines and dashes of purple-brown; columella wide, brown with white pillar; penis blunt, with small glandular disc at base; pallial oviduct large, of single multispiral loop, with capsule gland; oviparous. Tropical West Africa. Mangroves, driftwood, sheltered rocks............................................................................. Littoraria (Littoraria) cingulifera ( Dunker, 1845) ( Reid 1986: figs 4g, 99h, i)

- To 33 mm; 48–75 grooves on last whorl; colour cream, yellow or orange, with small brown dashes; columella narrow, purple to brown; penis bifurcate, glandular disc on projection of base, large smooth pointed filament; pallial oviduct small, without capsule gland, ovoviviparous (embryos in mantle cavity). Tropical West Africa. Mangroves, sheltered rocks.................................................. Littoraria (Littorinopsis) angulifera ( Lamarck, 1822) ( Reid 1986: figs 4o, 99d–f)

10 Globular to turbinate; irregular spiral ribs and threads unless eroded; colour variable; penis with at least 3 mamilliform glands, no glandular disc; pallial oviduct with insignificant spiral part, large septate brood pouch with embryos. NW Europe, Venice, Tunisia, Morocco, Canary Is, Azores, Namibia, South Africa. Rocky shores and lagoons.................................................................. Littorina (Neritrema) saxatilis ( Olivi, 1792) ( Reid 1996: figs 102–104, 106, 107)

- Penis with 1 mamilliform gland......................................................................... 11

11 Up to 27 fine threads on last whorl; brown to black with indistinct pattern of fine pale speckling; penis with 1 mamilliform gland, no glandular disc; pallial oviduct with 3 successive loops. Namibia, South Africa............................................................ Afrolittorina knysnaensis (Krauss in Philippi, 1847) ( Reid & Williams 2004: figs 19, 20)

- Penis with 1 mamilliform gland and glandular disc; pallial oviduct with albumen and capsule glands in 1 spiral.......... 12

12 12–20 low unequal ribs on last whorl; grey with brown or speckled ribs; St Helena....striate form of E. helenae ( Figs 22, 23)

- Regular spiral grooves or smooth; no pseudumbilicus; strong pattern of pale spots or lines on brown to black background.. 13

13 22–43 grooves above periphery of last whorl; 2 pale bands in aperture; penis with very large mamilliform gland occupying most of base, small glandular disc; first loop of albumen gland large; islands in Gulf of Guinea... E. soroziczac ( Figs 14, 15)

- 9–19 grooves above periphery of last whorl; 1 pale band in aperture; penis with small mamilliform gland and glandular disc; first loop of albumen gland small........................................................................ 14

14 Tall; 9–14 grooves above periphery of last whorl or smooth; Cape Verde Is................. E. caboverdensis ( Figs 9, 10)

- High turbinate; 9-19 grooves above periphery of last whorl; African mainland.................................... 15

15 Mediterranean to The Gambia, Canary Is.................................................. E. punctata ( Figs 6, 7)

- Sierra Leone to N Namibia.......................................................... E. pulchella ( Figs 12, 13)