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Published records, original data from recent field work on all of the islands of the Azores (NE Atlantic), and a revision of the entire mollusc collection deposited in the Department of Biology of the University of the Azores (DBUA) were used to compile a checklist of the shallow-water Polyplacophora of the Azores. Lepidochitona cf. canariensis and Tonicella rubra are reported for the first time for this archipelago, increasing the recorded Azorean fauna to seven species.
Azores, Mollusca, Polyplacophora, biodiversity, checklist
The marine molluscs of the Archipelago of the Azores are probably the best studied marine invertebrate group from these Atlantic Islands. Several taxonomic, ecological, genetic, biogeographic and recent palaeontological studies have greatly improved our knowledge of this phylum (see
Polyplacophorans (or “chitons”) include over 900 extant species worldwide that mostly live in shallow waters, usually on rocky substrates. They are oval in shape and dorso-ventrally flattened, neither tentacles nor eyes are present in the head region, and they possess eight distinctive overlapping shell plates or valves located on the dorsal side. These longitudinally arranged valves are surrounded by a muscular girdle, and the girdle covering or perinotum is ornamented with scales, spicules, bristles or other protuberances (
The present work is based on a review of new material collected from all the islands of the Azores, and updates the published information, documenting for the first time the occurrence of the polyplacophoran molluscs Lepidochitona cf. canariensis (Thiele, 1909) and Tonicella rubra (Linnaeus, 1767) in these oceanic islands.
A bibliographic review of the polyplacophoran species reported from the Atlantic Ocean was assembled. More than 1, 060 lots from the marine mollusc reference collection of the Department of Biology of the University of the Azores (DBUA – São Miguel Island), corresponding to approx. 850 dives in all islands, were examined and the polyplacophorans sorted and identified from 63 lots (303 specimens and 5 valves; see Table 1). The mollusc collection of the Department of Oceanography and Fisheries of the University of the Azores (DOP/ML – Faial Island) was also surveyed for chitons. Specimens were studied using either a Nikon SMZ 1000 or a Leica M125 stereomicroscope with incandescent light sources (Volpi Intralux 4100) and digital camera attached, which fed images to a desktop computer. The taxonomic organization of species in this list follows the morphological systematics of
Number of the sampling sites in the Reference Collection of the Department of Biology of the University of the Azores (DBUA) that yielded polyplacophorans, island and location of the sampling sites, depth range (m), number of Specimens (N. spc.), number of valves (N. valv.) and date.
DBUA | Island / seamount | Location | Depth range | N. spc. | N. valv. | Date |
---|---|---|---|---|---|---|
176 | São Miguel | Ponta da Pirâmide | 13 m | 1 | - | July-1988 |
190 | Flores | Fajã Grande | intertidal zone | 3 | - | 10-July-1989 |
191 | Flores | Ponta Delgada | intertidal zone | 1 | - | July-1989 |
193 | Flores | Santa Cruz (pool) | intertidal zone | 1 | - | 09-July-1989 |
240 | Flores | Santa Cruz | 20 m | 2 | - | July-1989 |
332 | Formigas Islets | West Bay | 6–8 m | 1 | - | 08-June-1990 |
337 | Formigas Islets | Formigas | intertidal zone | 7 | - | 06-June-1990 |
355 | Formigas Islets | Formigas | 15 m | - | 1 | 03-July-1991 |
356 | Formigas Islets | Formigas | intertidal zone | 1 | - | 01-July-1991 |
410 | Faial | Ilhéu Negro | 10 m | 1 | - | 24-July-1989 |
433 | Faial | Baía de entre-os-montes | ? | 1 | - | 26-July-1989 |
457 | Pico | Lajes do Pico | intertidal zone | 1 | - | July-1989 |
458 | Pico | Lajes do Pico | intertidal zone | 2 | - | July-1989 |
459 | Pico | Lajes do Pico | intertidal zone | 1 | - | July-1989 |
461 | Pico | Lajes do Pico | intertidal zone | 1 | - | July-1989 |
465 | Pico | Lajes do Pico | intertidal zone | - | 2 | July-1989 |
475 | Pico | Lajes do Pico | intertidal zone | - | 1 | July-1989 |
486 | Pico | Lajes do Pico | 2–4 m | 1 | - | July-1989 |
524 | Flores | Baixa da Calheta | intertidal zone | 1 | - | 28-Oct-1990 |
551 | Flores | Porto da Baleia | ? | 1 | - | 29-Oct-1990 |
554 | Flores | Pontinhas | intertidal zone | 6 | - | 28-Oct-1990 |
562 | Flores | Pontinhas | intertidal zone | 17 | - | 28-Oct-1990 |
569 | Flores | Baixa do Porto | ? | 1 | - | 28-Oct-1990 |
574 | Flores | Baixa do Porto | 6–12 m | 1 | - | 27-Oct-1990 |
577 | Flores | Lajes das Flores | 0–10 m | 2 | - | 27-Oct-1990 |
625 | São Miguel | São Roque | ? | 2 | - | 31-May-1991 |
637 | São Miguel | Ponta da Galera | 12 m | 1 | - | July-1989 |
662 | Pico | Lajes do Pico | 0–3 m | 3 | - | 19-Aug-1995 |
667 | Pico | Lajes do Pico | 0–6 m | 5 | - | 05-Aug-1996 |
671 | Pico | Lajes do Pico | intertidal zone | 1 | - | 18-Aug-1997 |
683 | São Miguel | São Vicente, Capelas | 10 m | 1 | - | 02-July-1996 |
700 | São Miguel | São Vicente, Capelas | 0–10.4 m | 2 | - | 17-July-1996 |
708/F | São Miguel | São Vicente, Capelas | 8 m | 1 | - | 19-July-1996 |
715 | São Miguel | Cerco, Caloura | intertidal zone | 3 | - | 26-Jan-1996 |
719 | São Miguel | Vila Franca Islet | 18 m | - | 1 | 03-Mar-1997 |
721 | São Miguel | Rosto do Cão Islet | 15 m | 1 | - | 15-May-1998 |
725 | Flores | Angra do Heroísmo bay | intertidal zone | 1 | - | 15-June-1998 |
730 | São Miguel | Baía da Pranchinha | 8.6 m | 3 | - | 04-July-1990 |
731 | São Miguel | Baía da Pranchinha | 13.8 m | 1 | - | 04-July-1990 |
732 | São Miguel | Baía da Pranchinha | 13.8 m | 5 | - | 04-July-1990 |
733 | São Miguel | Baía do Rosto do Cão | 9.5 m | 1 | - | 04-July-1990 |
740 | São Miguel | Baía do Rosto do Cão | intertidal zone | 28 | - | 26-June-1990 |
741 | São Miguel | Baía do Rosto do Cão | intertidal zone | 8 | - | 06-July-1990 |
743 | São Miguel | Baía do Rosto do Cão | intertidal zone | 1 | - | 06-July-1990 |
744 | São Miguel | Baía do Rosto do Cão | intertidal zone | 87 | - | 06-July-1990 |
745 | São Miguel | Baía do Rosto do Cão | intertidal zone | 53 | - | 06-July-1990 |
746 | São Miguel | Baía do Rosto do Cão | intertidal zone | 16 | - | 06-July-1990 |
747 | São Miguel | Baía do Rosto do Cão | intertidal zone | 2 | - | 06-July-1990 |
748 | São Miguel | Capelas | 14 m | 2 | - | 07-Oct-1996 |
751 | São Miguel | Baía do Rosto do Cão | subtidal | 1 | - | 05-May-1994 |
752 | São Miguel | Fenais da Luz | intertidal zone | 2 | - | 15-Apr-1992 |
767 | São Miguel | São Vicente, Capelas | 20.3 m | 1 | - | 11-July-1997 |
793 | São Miguel | São Vicente, Capelas | intertidal zone | 1 | - | 19-July-1997 |
794 | São Miguel | Ponta Delgada | ? | 2 | - | 20-Nov-1996 |
799 | Flores | Santa Cruz | tide pool | 1 | - | July-1999 |
800 | Pico | São João | 15 m | 1 | - | July-1999 |
801 | Faial | Monte da Guia | 23 m | 1 | - | 23-Aug-1999 |
803 | Faial | ? | 3–6 m | 3 | - | Sep-1998 |
857 | Pico | Monte | 8–10 m | 1 | - | 05-Aug-2005 |
858 | Pico | Barca, Madalena | 10 m | 1 | - | 09-Aug-2005 |
891 | D. João de Castro | seamount | 20 m | 1 | - | 31-Aug-2004 |
1047 | Pico | Ribeiras harbour | 4 m | 1 | - | 07-Aug-2000 |
1056 | São Miguel | Rosto do Cão Islet | intertidal zone | 4 | - | 26-Mar-2012 |
Animals ovate to elongate. Sculpture of the tegmentum varying from almost smooth to granular. Spicules and longer spines are present in the perinotum (
Tegmentum granulated. Overall, with the characteristics of the family.
http://species-id.net/wiki/Hanleya_hanleyi
Figs 1–2From the Barents Sea south to Algarve (Portugal), the Mediterranean Sea, Azores, Madeira, Canary Islands (
Flores (Porto da Baleia: DBUA 551, 1 spm), Pico (Lajes do Pico, 0–3m: DBUA 662, 1 spm).
No fossil representatives are known from the Azores.
Small (up to 22 × 13 mm), elongate oval, dorsal elevation ratio (intermediate valve height / valve width) up to ~0.3. Valves thick, not beaked and girdle narrow, with spicules. Intermediate valves rectangular. Tegmentum uniformly creamy white to light tan, occasionally with brown mineral deposits; uniformly sculpted with numerous large granules, arranged randomly on the lateral areas of the intermediate valves. Jugal (central) area distinct, with fine longitudinal riblets larger and more widely spaced than those in the pleural (outer) areas. Girdle perinotum covered in randomly distributed projecting spicules.
This species is very rare in the Azores.
Small to large in size [up to 110 × 80 mm – Eudoxochiton nobilis (Gray, 1843)], oval, tegmentum with fine or no apparent granular sculpture, valves appear smooth but with neat rows of black, pigmented shell eyes. Terminal valves multi-slitted (large numbers of insertion teeth), intermediate valves with 1–4 slits on insertion plates. Perinotum with small spicules.
Small to medium size [up to 55 × 36 mm – Callochiton dentatus (Spengler, 1797)]; extra-pigmentary eyes present. Overall, with the characteristics of the family.
http://species-id.net/wiki/Callochiton_septemvalvis
All Atlantic coasts of Europe, from Scandinavia, Britain and Ireland, south to the Mediterranean Sea, Morocco (
No material seen.
No fossil representatives are known from the Azores.
Moderate size (up to 22 × 14 mm), dorsal elevation ratio = 0.35 to 0.46, oval, valves beaked. Tegmentum very finely granulose, orange to brick red, often with white markings, or with shades of green, bright yellow, or bright orange. Intermediate valves rectangular. Sculpture smooth and glossy to the naked eye, diagonally set with black dots (the pigment cups of the ‘shell-eyes’); under magnification the valves are sculptured with small granules arranged in quincunx. Wide girdle, usually about 1/3 of the animal’s total width and covered in spicules, with a short marginal fringe of spicules. The girdle is coloured yellow or orange with red markings.
If this species does occur in the Azores, it must be very rare, as not a single specimen was found in the DBUA or DOP/ML collections. The species Callochiton septemvalvis was originally described from an abnormal specimen with seven valves.
Small to medium in size [up to 55 × 36 mm – Tonicella insignis (Reeve, 1847)], oval to elongate oval. Valve tegmentum appears smooth or granulose but without separated sculpture elements, terminal valves with multi-slitted insertion plates, intermediate valves with usually one insertion slit on each side. Girdle perinotum covered in small spicules.
Oval to elongate-oval, valve sculpture smooth to uniformly granular, girdle perinotum with irregular granules, most species with a short marginal fringe of blunt spicules.
This is the first record for the Azores.
The Mediterranean Sea (
Formigas Islets (intertidal zone: DBUA 337, 2 spm; DBUA 356, 1 spm), São Miguel (intertidal: DBUA 747, 1 spm).
No fossil representatives are known from the Azores.
Small size (up to 8.5 × 5 mm), dorsal elevation ratio = 0.39 (
This is a rare species, known only from a small number of specimens in the Azores.
Shallow polyplacophorans from the Azores. 1–2 Hanleya hanleyi (Bean in Thorpe, 1844), DBUA 551 (Flores, Porto da Baleia) 3–4 Lepidochitona cf. canariensis (Thiele, 1909), DBUA 356 (Formigas Islets, intertidal) 5 Lepidochitona piceola (Shuttleworth, 1853), DBUA 743 (São Miguel Island, Baía do Rosto do Cão, intertidal) 6–7 Lepidochitona simrothi (Thiele, 1902), DBUA 459 (Pico, Lajes do Pico, intertidal) 8 Tonicella rubra (Linnaeus, 1767), DBUA 891 (D. João de Castro seamount, 20 m depth) 9 Acanthochitona fascicularis (Linnaeus, 1767), DBUA 667 (Pico, Lajes do Pico, 0–6 m depth).
http://species-id.net/wiki/Lepidochitona_piceola
Fig. 5Azores and Canary Islands (
São Miguel (intertidal zone down to 14 m depth: DBUA 625, 1 spm; DBUA 743, 1 spm; DBUA 744, 23 spm; DBUA 745, 14 spm; DBUA 1056, 1 spm).
No fossil representatives are known from the Azores.
Small size (up to 10 × 6 mm), dorsal elevation ratio = 0.31 (
This species has been overlooked and confused with Lepidochitona simrothi. However, it is easy to separate these two species, as Lepidochitona piceola does not have the long, smooth, curved needles characteristic of the girdle of Lepidochitona simrothi. The only records before this work were those of
Azores (
Faial (3–6 m depth: DBUA 803, 3 spm), Flores (all samples collected in the intertidal zone: DBUA 190, 3 spm; DBUA 191, 1 spm; DBUA 193, 1 spm; DBUA 524, 1 spm; DBUA 554, 6 spm; DBUA 562, 17 spm), Formigas Islets (intertidal zone: DBUA 337, 5 spm; DOP/ML 0032, 1 spm), Pico (0–3 m depth: DBUA 457, 1 spm; DBUA 458, 2 spm; DBUA 459, 1 spm; DBUA 461, 1 spm; DBUA 465, 2 valves; DBUA 475, 1 valve; DBUA 662, 2 spm); São Miguel (intertidal zone down to 14 m depth: DBUA 625, 1 spm; DBUA 715, 3 spm; DBUA 732, 3 spm; DBUA 740, 28 spm; DBUA 741, 8 spm; DBUA 744, 64 spm; DBUA 745, 39 spm; DBUA 746, 16 spm; DBUA 747, 1 spm; DBUA 793, 1 spm).
No fossil representatives are known from the Azores.
Animal rather small (up to 8 × 4 mm), dorsal elevation ratio = 0.37 (
Specimens recorded by
With the characteristics of the family. Valves with ‘spongy’ eaves (i.e. porous, penetrated laterally by large aesthete canals).
http://species-id.net/wiki/Tonicella_rubra
Fig. 8This is the first record for the Azores.
This species has an Arctic-circumboreal distribution, including the Arctic Ocean (Barents Sea, White Sea, Spitzbergen), the North Pacific (northern Japan), the western North Atlantic as far south as New London (Connecticut) (
Dom João de Castro seamount (20 m depth: DBUA 891, 1 spm).
No fossil representatives are known from the Azores.
Up to 15 × 9 mm in the North Atlantic; dorsal elevation ratio = 0.29. Valves beaked, girdle narrow. Tegmentum appears smooth, with growth lines clearly visible under magnification. Colour orange to pinkish, generally with small reddish-brown blotches. Girdle relatively narrow and covered by small scales, appearing sandy to the naked eye, coloured like the tegmentum, but sometimes with white or cream markings particularly at the junctures between valves.
The Dom João de Castro Bank (Lat 38°13.3’N, Long 26°36.2’W) is a shallow seamount (minimum depth = 13 m) located between the islands of São Miguel and Terceira. The last eruption was in December 1720 when a small island (~1 km long and 150 m high) was formed (
The broad girdle is covered by coarse spines and partially covers the valves. The valve tegmentum (dorsal aspect) is reduced relative to the articulamentum (ventral part covered by the girdle). Head valve usually with five slits in margin.
Girdle with large distinct bristles, clumps of long straight spicules, emerging from the girdle at the junctures between the valves on both sides.
http://species-id.net/wiki/Acanthochitona_fascicularis
Fig. 9Found in the North Atlantic from Ireland and Britain, south to Portuguese shores (
Faial (10–23 m depth: DBUA 410, 1 spm; DBUA 433, 1 spm; DBUA 801, 1 spm), Flores (intertidal zone down to 20 m depth: DBUA 240, 2 spm; DBUA 569, 1 spm; DBUA 574, 1 spm; DBUA 577, 2 spm; DBUA 725, 1 spm; DBUA 799, 1 spm), Formigas (6–15 m depth: DBUA 332, 1 spm; DBUA 355, 1 valve), Pico (intertidal zone down to 15 m depth: DBUA 486, 1 spm; DBUA 667, 5 spm; DBUA 671, 1 spm; DBUA 800, 1 spm; DBUA 857/DOP/ML 0050, 1 spm; DBUA 858/DOP/ML0051, 1 spm; DBUA 1047, 1 spm), São Miguel (intertidal zone down to 20 m depth: DBUA 176, 1 spm; DBUA 637, 1 spm; DBUA 683, 1 spm; DBUA 700, 2 spm; DBUA 708/F, 1 spm; DBUA 719, 1 valve; DBUA 721, 1 spm; DBUA 730, 3 spm; DBUA 731, 1 spm; DBUA 732, 2 spm; DBUA 733, 1 spm; DBUA 748, 2 spm; DBUA 751, 1 spm; DBUA 752, 2 spm; DBUA 767, 1 spm; DBUA 794, 2 spm; DBUA 1056, 3 spm).
No fossil representatives are known from the Azores.
Rather large, up to 24 × 15 mm (DBUA 667). Overall colour is variable (black, blue, olive, orange, cream). Valves typically olive-green with reddish blotches; the wide jugal area is usually lighter (cream or beige) with dark longitudinal streaks. Visible portion of the valves is trapezoidal, little or no beak. The central area with faint longitudinal grooves; lateral areas covered with closely-spaced, small round granules in quincunx and forming radiating rows. Girdle broad and densely covered with short spicules, with 18 large tufts of spines arranged around the head and at the sutures. One specimen from the Azores had a girdle that appeared cyan blue in life. There is a distinct marginal fringe of small tapered spicules.
This is the largest chiton in the Azores, commonly found under stones buried in pebbles or in sand, in shallow water, sometimes in groups of two or three specimens. Acanthochitona fascicularis is particularly variable in colour.
We thank the Direcção Regional da Ciência, Tecnologia e Comunicações (Regional Government of the Azores), and FCT (Fundação para a Ciência e a Tecnologia) of the Portuguese government for financial support. Helpful comments from Bruce A. Marshall and Bruno Dell’Angelo improved this manuscript. Sérgio Ávila acknowledges his Ciência 2009 research contract funded by FCT.