Peneroplis de Monfort 1808

Mamo, Briony L., 2016, Benthic Foraminifera from the Capricorn Group, Great Barrier Reef, Australia, Zootaxa 4215 (1), pp. 1-123 : 62-65

Peneroplis de Monfort 1808

Peneroplis pertusus ( Forskål 1775) ( Fig. 14 :17–20; Fig. 15 :1–5)

1775 Nautilus pertusus Forskål , p. 125, nomen nudum.

1884 Peneroplis pertusus (Forskål) ; Brady, pl. 13, fig. 16, 17.

1921a Peneroplis pertusus (Forskål) ; Cushman, p. 75, pl. 18, figs 7, 8. 1978 Peneroplis pertusus (Forskål) ; Cheng & Zheng, p. 192, pl. 14, figs 8, 9. 1984 Peneroplis pertusus (Forskål) ; Hallock, p. 251, fig. 1: 3.

1991 Peneroplis pertusus (Forskål) ; Van Marle, p. 72, pl. 4, fig. 10.

1992b Peneroplis pertusus (Forskål) ; Hatta & Ujiié, p. 78, pl. 16, fig. 1. 1993 Peneroplis planatus (Fichtel & Moll) ; Hottinger et al., p. 70, pl. 79, figs 2, 5, 13, 15. 1994 Peneroplis pertusus (Forskål) ; Gudmundsson, p. 115, text figs 23, 24, pl. 3, figs 1, 3. 1994 Peneroplis pertusus (Forskål) ; Loeblich & Tappan, p. 62, pl. 110, figs 1–5. 1995 Peneroplis pertusus (Forskål) ; Lobegeier, p. 75, pl. 3, figs 14–16. 1999 Peneroplis spp.; Harney et al., p. 64, fig. 1.

1999 Peneroplis pertusus (Forskål) ; Hohenegger et al., p. 128, fig. 9. 2002 Peneroplis pertusus (Forskål) ; Bicchi et al., fig. 7:6.

2002 Peneroplis pertusus (Forskål) ; Yordanova & Hohenegger, p. 182, pl. 29, figs 10–13. 2005 Peneroplis pertusus (Forskål) ; Goldbeck et al., pl. 1, fig. 5.

2009 Peneroplis pertusus (Forskål) ; Parker, p. 152, figs 108a–l.

2010 Peneroplis pertusus (Forskål) ; Narayan & Pandolfi, p. 2076, pl. 1, fig. 29. 2012 Peneroplis pertusus (Forskål) ; Debenay, p. 113, pl. 4.

Description. See Gudmundsson (1994, p. 115, text figs 23, 24, pl. 3, figs 1, 3), Hohenegger et al. (1999, p. 128, fig. 9), Yordanova & Hohenegger (2002, p. 182, pl. 29, figs 10–13) and Parker (2009, p. 152, figs 108a–l, 109a–l).

Remarks. Peneroplis pertusus ( Forskål 1775) is a well-known species that consistently displays variation in morphology that commonly makes confident specific assignment difficult ( Boltovskoy & Wright 1976). Forskål (1775) never illustrated nor elected a holotype with the original description of this species as Nautilus pertusus and therefore, similar to Van Marle (1991) is considered nomen nudum. Attempts have been made to subdivide this species ( Heron-Allen & Earland 1915; Gudmundsson 1994), but a definitive classification for this taxon is still elusive.

Specimens from the CG were assigned to P. pertusus due to their compressed planispiral test with an adult stage that can uncoil, fan out and in some cases become partially uniserial. Apertures are terminal, but the size, shape, arrangement and number can differ greatly within this species ( Fig. 14 :17–20; Fig. 15 :1–5). Juvenile specimens bear varying arrangements of crenulated apertures that can divide at the base of the apertural face. During ontogeny, the apertures can also take the form of sub-parallel vermicular slits that become more parallel and uniform in nature the closer the specimens get to the fanning, uniserial ontogenetic stage. The chamber walls are ornamented by numerous low costae that follow the direction of coiling and are interrupted by depressed sutures.

The juvenile form of P. pertusus is similar to Peneroplis antillarum d’Orbigny 1839 whilst adult forms closely resemble Peneroplis planatus (Fichtel & Moll 1993) . However, as outlined by Hohenegger et al. (1999), P. pertusus can be differentiated from both P. antillarum by possessing evolute (as opposed to involute) planispiral enrolment in later chambers and from P. planatus by lacking fanning outer chambers in combination with a pronounced umbilicus. Additionally, the sutures of P. pertusus are more strongly depressed and the final section of adult specimens never display flaring chambers. However, it is now clear that some adult P. pertusus do in fact have flaring final chambers ( Brady 1884; Loeblich & Tappan 1994; Saraswati et al. 2004; Goldbeck et al. 2005; Parker 2009).

Forskål (1775) never illustrated nor elected a holotype with the original description of this species and there exists substantial morphological variety. The P. pertusus specimens from Shell Cove collected by Brady (1884) exhibit either a stout, lenticular, biconvex morphology or a flat, large, flaring test shape. Further morphological variation includes types with a broad septal margin and thin, deeply depressed sutures that give the test a lobate outline. Morphotypes with a large, flaring test have thin, weakly depressed sutures that gave the test a smooth or weakly lobate outline. Specimens assigned to P. planatus from Wood Lark Island also display these morphological variants, however, most specimens have the large, flat, flaring test morphology associated with P. planatus . Furthermore , the central axis from which the sutures flare radially is not centered in these specimens but is offset to one side creating an asymetrical shape. Peneroplis pertusus specimens from Shell Cove possessed this feature as well, but it is not as exaggerated. Strong morphological similarities under both taxonimic names exist and other authors have combined both morphotyptes under P. pertusus ( Cheng & Zheng 1978; Parker 2009) or P. planatus ( Hottinger et al. 1993) however it is the large, flat and flaring test morphology and linear arrangement of regular to irregular apertures that establishes P. planatus from P. pertusus . Due to the large amount of literature that separates these two species based on these morphological characteristic, CG specimens have been similarly separated.

Peneroplis pertusus has a cosmopolitan distribution. Originally collected from the Suez Canal, Egypt ( Forskål 1775), P. pertsus has been collected extensively throughout the Pacific from Australia ( Brady 1884; Gudmundsson 1994; Lobegeier 1995; Parker 2009; Narayan & Pandolfi 2010) at depths of 5–46 m, north to Indonesia and the South-west Pacific Islands ( Van Marle 1991; Loeblich & Tappan 1994) at depths down to 132 m, from Japan ( Hatta & Ujiié 1992b; Hohenegger et al. 1999; Yordanova & Hohenegger 2002; Saraswati et al. 2004), China ( Cheng & Zheng 1978) and Central Pacific ( Hallock 1984; Harney et al. 1999; Bicchi et al. 2002). It has additionally been found throughout the Carribean ( Cushman 1921a; Gudmundsson 1994), the Meditteranean ( Goldbeck et al. 2005) and Egypt ( Hottinger et al. 1993).

Distribution within study area. Peneroplis pertusus occurred in all sampled CG reefs with an average number abundance of approximately five specimens per site. However, this species was by far the most abundant on the reef rampart at site 10 of the ST/HW transect across Heron Reef flat where 94 specimens were counted. Other notable occurrences were also recorded from sites 50 and 52 in One Tree Lagoon 2, with 29 and 20 specimens collected, respectively. The lowest abundance of P. pertusus was recorded in Heron Lagoon where only eight specimens were collected from three sites.

Peneroplis planatus ( Fichtel & Moll 1798) ( Fig. 15 :6–7)

1798 Nautilus planatus var. α Fichtel & Moll, p. 91, pl. 16, figs a–c, i.

1798 Nautilus planatus var. β Fichtel & Moll, p. 93, pl. 16, figs d, e.

1798 Nautilus planatus var. γ Fichtel & Moll, p. 94, pl. 16, figs g, h.

1884 Peneroplis planatus (Fichtel & Moll) ; Brady, pl. 13, fig. 15.

1921a Peneroplis planatus (Fichtel & Moll) ; Cushman, p. 75, pl. 18, figs 9, 13. 1978 Peneroplis pertusus (Forskål) ; Cheng & Zheng, p. 192, pl. 14, figs, 11, 14. 1984 Peneroplis planatus (Fichtel & Moll) ; Hallock, p. 251, fig. 1: 2.

1988 Peneroplis planatus (Fichtel & Moll) ; Loeblich & Tappan, p. 371, pl. 391, figs 7, 8. 1991 Peneroplis planatus (Fichtel & Moll) ; Van Marle, p. 73, pl. 4, fig. 9.

1992b Peneroplis planatus (Fichtel & Moll) ; Hatta & Ujiié, p. 79, pl. 16, fig. 2. 1993 Peneroplis planatus (Fichtel & Moll) ; Hottinger et al., p. 70, pl. 80, figs 2–4. 1999 Peneroplis planatus (Fichtel & Moll) ; Hohenegger et al., p. 128, fig. 8. 2002 Peneroplis planatus (Fichtel & Moll) ; Yordanova & Hohenegger, p. 182, pl. 29, figs 6–9. 2009 Peneroplis pertusus (Forskål) ; Parker, p. 152, fig. 109a–l.

2012 Peneroplis planatus (Fichtel & Moll) ; Debenay, p. 114, pl. 4.

Description. See Hohenegger et al. (1999, p. 128, fig. 8), Yordanova & Hohenegger (2002, p. 182, pl. 29, figs 6–9) and Debenay (2012, p. 114, pl. 4).

Remarks. Despite lectotypes being later established and illustrated for this species ( Rögl & Hansen 1984), as previously commented for P. pertusus , there is significant difficulty in establishing P. planatus as a separate species when P. pertusus encompasses such a broad range of morphologies, incluing those resembling P. planatus . However, due to the significant amount of literature that does split P. pertusus and P. planatus , CG specimens of this species have been identified due to their flat cross-section, their flaring outer chambers ( Fig. 15 :6) and their long row of linearly arranged apertures ( Fig. 15 :7).

The most varying morphological features between specimens of P. plantus include the shape of the apertures and the flaring extent of the final chambers. Different taxonomic descriptions and manuscripts have described the apertures of P. planatus as both regular ( Hohenegger et al. 1999) and irregular ( Debenay 2012). Capricorn Group specimens possess a regular aperture shape ( Fig. 15 :3) bearing strong similarity to others specimens of the Pacific ( Brady 1884; Cushman 1921a; Hallock 1984; Hohenegger et al. 1999).

Fichtel & Moll’s (1798) original specimens were reported from Tuscany and possesses a global distribution ( Admiralty Islands from 29–46 m—Brady 1884; Monego Bay , Jamaica—Cushman 1921a; Xisha Islands—Cheng & Zheng 1978; Palau and Oahu Reefs—Hallock 1984; eastern Indonesia—Van Marle 1991; Mediterranean— Loeblich & Tappan 1988; Gulf of Aqaba lower photic zone—Hottinger 1993; Ryukyu Island Arc—Hatta & Ujiié 1992b, Hohenegger et al. 1999 and Yordanova & Hohenegger 2002; Ningaloo Reef—Parker, 2009; New Caledonia southwestern lagoon down to 40 m—Debenay 2012).

Distribution within study area. Peneroplis planatus was found in low numbers (never more than 8 specimens per site) within Heron, One Tree and Wistari Reefs within the CG. It was most commonly collected from the Heron Reef flat and scarce within Wistari Reef where it was collected only once from a single site (24).

Figures

FIGURE 14. (Unless otherwise specified all scale bars = 100 µm). 1 – 4. Pseudohauerina involuta (Cushman, 1946). 1 – 2. QM # G 465873 from CHANNEL, lateral (1) and angled lateral (2) views. 3 – 4. QM # G 465874 from CHANNEL, lateral (3) and oral (4) views. Scale bars = 200 µm. 5 – 10. Alveolinella quoyi (d’Orbigny, 1826). 5 – 7. QM # G 465965 from CHANNEL, oral (5), lateral (6) and peripheral (7) views. 8 – 9. QM # G 465964 from CHANNEL, oral (8) and lateral (9) views. Scale bar for 8 = 200 µm. 10. QM # G 465865 from CHANNEL, lateral view, scale bar = 500 µm. 11 – 16. Monalysidium acicularis (Batsch, 1791). 11 – 13. QM # G 465778 from ST / HW 2, lateral view (11, 12) and oral (13) views. Scale bar = 200 µm. 14 – 16. QM # G 465888 from ST / HW 4 lateral (14, 15) and oral (16) views. 17 – 20. Peneroplis pertusus (Forskål, 1775). 17 – 18. QM # G 465882 from ST / HW 2, lateral (17) and oral (18) views. 19 – 20. QM # G 465804 from T 2 – 3, lateral (19) and oral (20) views.

FIGURE 15. (Unless otherwise specified all scale bars = 100 µm). 1 – 5. Peneroplis pertusus (Forskål, 1775). 1 – 3. QM # G 465896 from ST / HW 10, lateral (1, 2) and oral (3) views. 4 – 5. QM # G 465805 from T 2 – 3, lateral (4) and oral (5) views. 6 – 7. Peneroplis planatus (Fichtel & Moll, 1798) QM # G 465995 from T 1 – 10, lateral (6) and oral (7) views. Scale bar = 500 µm. 8 – 13. Parasorites sp. 1. 8 – 9. QM # G 465802 from T 2 – 3, lateral (8) and oral (9) views. Scale bar = 200 µm. 10. QM # G 465781 from ST / HW 2, lateral view. Scale bar = 200 µm. 11 – 13. QM # G 465982 from OTI 54, lateral (11), close up of intradermal plates (12) and oral (13) views. Scale bar for 12 = 50 µm. 14 – 20. Marginopora vertebralis Quoy & Gaimard, 1830. 14 – 16. QM # G 465793 from T 2 – 1, lateral view (14), scale bar = 200 µm. 15. Oral view. 16. Close-up oral view, scale bar = 50 µm. 17 – 18. QM # G 466030 from T 2 – 5, lateral (17) and close-up oral (18) views. Scale bar for 17 = 1 mm. 19 – 20. QM # G 466020 from T 1 – 2, lateral (19), oral (20) views. Scale bar for 19 = 2 mm. Scale bar for 20 = 1 mm.