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Ossicle micrographs of collected specimens. A, Euapta godeffroyi ( uf 11413): miliary granules from body wall; B, E. godeffroyi ( uf 11413): anchor from body wall; C, E. godeffroyi ( uf 11413): anchor plate from body wall; D, Opheodesoma grisea ( uf 5855): miliary granules from body wall; E, O. grisea ( uf 5855): anchor from body wall; F, O. grisea ( uf 5855): anchor plate from body wall; G, Polyplectana sp. ( uf 5874): anchor from body wall; H, Polyplectana sp. ( uf 5874): anchor plate from body wall. Scale indicates 50 μ m for A and D, and 100 μ m for B, C, E, F, G, H . 

Ossicle micrographs of collected specimens. A, Euapta godeffroyi ( uf 11413): miliary granules from body wall; B, E. godeffroyi ( uf 11413): anchor from body wall; C, E. godeffroyi ( uf 11413): anchor plate from body wall; D, Opheodesoma grisea ( uf 5855): miliary granules from body wall; E, O. grisea ( uf 5855): anchor from body wall; F, O. grisea ( uf 5855): anchor plate from body wall; G, Polyplectana sp. ( uf 5874): anchor from body wall; H, Polyplectana sp. ( uf 5874): anchor plate from body wall. Scale indicates 50 μ m for A and D, and 100 μ m for B, C, E, F, G, H . 

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In December 2002, July 2007 and December 2009, we surveyed the sea cucumber fauna of the western Caroline Island of Yap (Federated States of Micronesia). We collected 37 species of holothuroids, including 32 species of aspidochirotes and five species of apodans. We found all 13 of the previously reported species and 24 new records for the islands—1...

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Context 1
... 1952:23, pl. 3, fig. 4, text fig. 8 a–t; Rowe and Gates, 1995:325; Samyn, 2000:15. Stichopus cf. monotuberculatus: Samyn and Vanden Berghe, 2000:31, pl. 2 F–H . material examined : uf 11415, Kaday, < 1 m depth during the day under rubble on middle reef flat, 11 December 2009. remarks : This species is a new record for Yap. This nocturnal species displayed an ar- ray of color patterns, also seen in other loca- tions in the Indo-Pacific (Byrne et al. 2010, Michonneau et al. 2013). The background varies from beige to camel with green and dark brown mottling. This species was com- monly observed under rocks and coral heads during the day and exposed on sand or coral- line rock at night. Recent genetic work by Byrne et al. (2010) revealed that S. horrens and S. monotuberculatus share similar sets of vari- able color patterns. The two species are dis- tinguished by ossicle morphology: S. horrens possesses tacklike body wall ossicles, whereas S. monotuberculatus does not (Byrne et al. 2010). The collected specimen was devoid of tacklike ossicles (Figure 4 O, P, Q ); however, ubiquity of the shared color forms in Yap sug- gests that S. horrens may also inhabit Yapese reefs. Stichopus vastus Sluiter, 1887 Stichopus vastus Sluiter, 1887:198–199, pl. 2, figs. 46 – 48; Sluiter, 1895:79; Rowe and Gates, 1995:381–382; Forbes et al., 1999:14; Massin, 1999:71–77, figs. 57 a–l, 58 a–m, 59 a–g, 60 a–d, 61, 112 d–e . Stichopus sp.: Allen and Steene, 1996:246; Gosliner et al., 1996:282, fig. 1039. material examined : uf 5367, 5869, 5913, O’Keefe’s Island, < 1 m depth during the day in sea grass bed near patchy reef, 30 July 2007. remarks : This species is a new record for Yap. It was uncommon and seen during the day and night on silty sand near mangroves and in the inner harbor. Genus Thelenota H. L. Clark, 1921 Thelenota ananas ( Jaeger, 1833) Trepang ananas Jaeger, 1833:24, pl. 3 fig. 1; H. L. Clark, 1922:48. Holothuria ananas: Quoy & Gaimard, 1833:110, pl. 6, figs. 1–3. Holothuria ( Thelenota ) ananas: Brandt, 1835:53. Stichopus ananas: Semper, 1868:75; Ludwig, 1882:133; Sluiter, 1901:30, pl. 2, fig. 1; Mitsukuri, 1912:150 –155, pl. 1, figs. 6 –8, text fig. 25. Thelenota ananas: A. M. Clark and Rowe, 1971:178–179, fig. 87 a; Rowe and Doty, 1977:227, figs. 2 b, 5 e; Grosenbaugh, 1981:51–53; Cherbonnier, 1988:152–153, fig. 63 a–k; Rowe and Gates, 1995:382. material examined : uf 5857, Balabat, 5 m depth during the day on pavement, 2 August 2007. remarks : This survey suggests that T. ananas is uncommon in Yap, relative to other locations in the western Pacific. It was oc- casionally seen exposed during the day and night on the outer reef slope on coralline rock or sand to at least 20 m deep. Thelenota rubralineata Massin & Lane, 1991 Plate III C Thelenota rubralineata Massin & Lane, 1991:57– 64, figs. 1–8; Kerr et al. 2007:21– 22, fig. 4. material examined : Photo record (Plate III C ), Ulithi Atoll, 25 m depth. Picture from Vanessa Fread, 27 July 2007. remarks : This species is a new record for Yap. Several specimens of this unmistakable and striking species have been seen by Brian Greene ( University of Hawai‘i at Mänoa, Ho- nolulu, pers. comm., 2007) around Yap and other islands in Yap State, on the deep outer reef slopes to at least 60 m depth. Order Apodida Family Chiridotidae Genus Chiridota Eschscholtz, 1829 Chiridota sp. Plate III D material examined : Photo record (Plate III D ), site unrecorded, exposed near coral rubble on middle reef flat, 28 November 2002. remarks : This species is a new record for Yap. This species usually takes shelter under rubble, often near sandy areas. Chiridota sp. often cleaves its body transversely when disturbed, as shown in Plate III D . This is likely either C . rigida Semper, 1868, or C. hawaiiensis Fisher, 1907, two widespread and poorly characterized species in the Pacific (A. M. Clark and Rowe 1971, Paulay 2003). Family Synaptidae Genus Euapta Oestergren, 1898 Euapta godeffroyi (Semper, 1868) Figure 5 A, B, C Synapta godeffroyi Semper, 1868:231, pl. 39, fig. 13. Euapta godeffroyi: Oestergren, 1898:113; H. L. Clark, 1907:72; H. L. Clark, 1921:158; Cherbonnier, 1955:172, pl. 48, figs. a–j; A. M. Clark and Rowe, 1971:185–186, pl. 30, fig. 8; Rowe and Doty, 1977:235– 236, figs. 5 c, 8 h; Cherbonnier, 1988:249– 251, fig. 111 a–j; Kerr, 1994:171; Rowe and Gates, 1995:338; Rowe and Richmond, 2004:3307–3308. material examined : uf 11413, Gilfith, < 1 m depth at night on rubble on middle reef flat, 14 December 2009. remarks : This species is a new record for Yap. The few other specimens of Euapta seen in this survey also all appear to be this species, rather than the closely related E . tahitiensis Cherbonnier, 1955, which also occurs in Mi- cronesia (Michonneau et al. 2013). This spe- cies was active at night on rubble and sand. Specimens seen during the day were hidden under rocks and rubble. Genus Opheodesoma Fisher, 1907 Opheodesoma grisea (Semper, 1868) Plate III F, G, H; Figure 5 D, E, F Synapta grisea Semper, 1868:11, pl. 4, figs. 6, 7. Opheodesoma grisea: Fisher, 1907:723; Heding, 1928:129, figs. 4.7, 7.3, 9; Cherbonnier, 1955:171, pl. 49, figs. k–t; A. M. Clark and Rowe, 1971:186, pl. 30, fig. 11; Rowe and Doty, 1977:235, figs. 5 a, 8 g; Cherbonnier, 1988:244 –246, fig. 109 a–f; Rowe and Gates, 1995:340; Rowe and Richmond, 2004:3308. material examined : uf 5862, 5876, Fanif, 10 m in blue hole, 3 August 2007. uf 5885, 5886, O’Keefe’s Island, 1 m depth in sea grass near massive Porites sp. colonies, 30 July 2007. uf 5855, 5900, O’Keefe’s Island, 1 m depth in sea grass near mangroves, 5 August 2007. uf 11406, Kaday, < 1 m depth during the day in sea grass bed on inner reef flat, 11 December 2009. remarks : This species is a new record for Yap. We found diverse color patterns among specimens referable to O. grisea (Plate III F, G, H ). This species was frequently observed in sea grass on inner reef flats, as well as at night in areas with high coral cover in the blue holes. Genus Polyplectana H. L. Clark, 1907 Polyplectana sp. Figure 5 G, H material examined : uf 5874, Maap, 2 m depth during the day in sea grass bed on inner reef flat, 29 July 2007. remarks : This species is a new record for Yap. This is a small species found in large numbers in sea grass, bordering a patch reef of Acropora spp. Absence of miliary granules and branched ossicles suggests that this spe- cies is P. kerfersteini. Genus Synapta Eschscholtz, 1829 Synapta maculata (Chamisso & Eysenhardt, 1821) Plate III E Holothuria maculata Chamisso & Eysenhardt, 1821:352, pl. 25; H. L. Clark, 1907:78, pls. 1, 4, figs. 17–19, 26; H. L. Clark, 1921:160; A. M. Clark, 1952:204; Cherbonnier, 1955:170, pl. 47, figs. a–d; A. M. Clark and Rowe 1971:186 –187, pl. 30, fig. 9. Synapta maculata andreae: Heding, 1928:115, text fig. 3, fig. 2. Synapta maculata sundaensis: Heding, 1928:116, text fig. 3, figs. 3–5. Synapta maculate: Heding, 1928:113, fig. 2; Rowe and Doty, 1977:234 –235, figs. 5 a, 8 e; Cherbonnier, 1988:107–108, fig. 43 a– k; Kerr, 1994:171; Rowe and Gates, 1995:343–344; Rowe and Richmond, 2004:3308. material examined : Photo record (Plate III E ), Gagil, 1.5 m depth in grass flats mixed with corals, 26 November 2002. uf 5894, Balabat, 15 m on sandy bottom of channel, 2 August 2007. uf 11424, Kaday, < 1 m depth during the day in sea grass bed on inner reef flat, 10 December 2009. remarks : This species is a new record for Yap. This large, diurnally active species was commonly found in sea grass. Specimens were also found rarely in silty sites, scattered with occasional coral colonies. Despite the rich fauna, only a handful of accounts report the holothuroids of Micro- nesia. The most recent and comprehensive account from Guam (Michonneau et al. 2013) enumerated 64 species of sea cucumbers, in- cluding 18 new records. The 37 species re- corded in this study is comparable to the di- versity known from Guam before the recent update (46 species) and makes Yap the best- documented Micronesian island after Guam. The known species composition of Yap is similar to that of Guam; the aspidochirotes dominate the reefs and adjacent habitats in Yap (86%) and Guam (81%), and the dendro- chirotes are the most poorly reported group in Yap (0%) and Guam (5%). Of the 37 holothuroid species recorded in this study, 24 (19 aspidochirotes and five apodans) are new records. At least two spe- cies, Actinopyga sp. and Holothuria ( Theelothuria ) sp., appear to be undescribed. One of the species we collected, H. ( Semperothuria ) roseomaculata , was also undescribed at the time of the surveys but was recently described (Kerr 2013). A few of the species were re ported only as photo records because we lacked permis- sion to collect during one visit. Two of these species, H . aff. ( Lessonothuria ) cavans and H . (? Thymiosycia ) sp., may represent rare species; however, we cannot make confident identifi- cations because we lack voucher specimens. The small-scale distribution patterns of holothuroids are closely related to micro- habitat and reef zonation (Levin 1979, Sloan 1979, 1982, Kerr et al. 1993, Kerr 1994). The distribution of Yapese holothuroids also varied considerably between reef zonations (Table 1). Sea grass beds on the inner reef flats were inhabited by the largest diversity of apodans, Opheodesoma grisea, Polyplectana sp., and Synapta maculata, as well as small cryptic aspidochirotes H . aff . ( Lessonothuria ) cavans and H . (? Thymiosycia ) sp., and the commer- cially valuable aspidochirotes Holothuria ( Metriatyla ) ? lessoni and H . ( Metriatyla ) scabra . The diurnally burying Bohadschia species, B. marmorata, B. vitiensis, and B. koellikeri, were also frequently observed in sea grass beds and near mangroves during the late afternoon and at night. Diurnally cryptic species, H. coluber, H. hilla, H. impatiens, H. pervicax, H. turriscelsa, Chiridota sp., Euapta godeffroyi, and Stichopus cf. monotuberculatus, were ...
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... O, P, Q ); however, ubiquity of the shared color forms in Yap sug- gests that S. horrens may also inhabit Yapese reefs. Stichopus vastus Sluiter, 1887 Stichopus vastus Sluiter, 1887:198–199, pl. 2, figs. 46 – 48; Sluiter, 1895:79; Rowe and Gates, 1995:381–382; Forbes et al., 1999:14; Massin, 1999:71–77, figs. 57 a–l, 58 a–m, 59 a–g, 60 a–d, 61, 112 d–e . Stichopus sp.: Allen and Steene, 1996:246; Gosliner et al., 1996:282, fig. 1039. material examined : uf 5367, 5869, 5913, O’Keefe’s Island, < 1 m depth during the day in sea grass bed near patchy reef, 30 July 2007. remarks : This species is a new record for Yap. It was uncommon and seen during the day and night on silty sand near mangroves and in the inner harbor. Genus Thelenota H. L. Clark, 1921 Thelenota ananas ( Jaeger, 1833) Trepang ananas Jaeger, 1833:24, pl. 3 fig. 1; H. L. Clark, 1922:48. Holothuria ananas: Quoy & Gaimard, 1833:110, pl. 6, figs. 1–3. Holothuria ( Thelenota ) ananas: Brandt, 1835:53. Stichopus ananas: Semper, 1868:75; Ludwig, 1882:133; Sluiter, 1901:30, pl. 2, fig. 1; Mitsukuri, 1912:150 –155, pl. 1, figs. 6 –8, text fig. 25. Thelenota ananas: A. M. Clark and Rowe, 1971:178–179, fig. 87 a; Rowe and Doty, 1977:227, figs. 2 b, 5 e; Grosenbaugh, 1981:51–53; Cherbonnier, 1988:152–153, fig. 63 a–k; Rowe and Gates, 1995:382. material examined : uf 5857, Balabat, 5 m depth during the day on pavement, 2 August 2007. remarks : This survey suggests that T. ananas is uncommon in Yap, relative to other locations in the western Pacific. It was oc- casionally seen exposed during the day and night on the outer reef slope on coralline rock or sand to at least 20 m deep. Thelenota rubralineata Massin & Lane, 1991 Plate III C Thelenota rubralineata Massin & Lane, 1991:57– 64, figs. 1–8; Kerr et al. 2007:21– 22, fig. 4. material examined : Photo record (Plate III C ), Ulithi Atoll, 25 m depth. Picture from Vanessa Fread, 27 July 2007. remarks : This species is a new record for Yap. Several specimens of this unmistakable and striking species have been seen by Brian Greene ( University of Hawai‘i at Mänoa, Ho- nolulu, pers. comm., 2007) around Yap and other islands in Yap State, on the deep outer reef slopes to at least 60 m depth. Order Apodida Family Chiridotidae Genus Chiridota Eschscholtz, 1829 Chiridota sp. Plate III D material examined : Photo record (Plate III D ), site unrecorded, exposed near coral rubble on middle reef flat, 28 November 2002. remarks : This species is a new record for Yap. This species usually takes shelter under rubble, often near sandy areas. Chiridota sp. often cleaves its body transversely when disturbed, as shown in Plate III D . This is likely either C . rigida Semper, 1868, or C. hawaiiensis Fisher, 1907, two widespread and poorly characterized species in the Pacific (A. M. Clark and Rowe 1971, Paulay 2003). Family Synaptidae Genus Euapta Oestergren, 1898 Euapta godeffroyi (Semper, 1868) Figure 5 A, B, C Synapta godeffroyi Semper, 1868:231, pl. 39, fig. 13. Euapta godeffroyi: Oestergren, 1898:113; H. L. Clark, 1907:72; H. L. Clark, 1921:158; Cherbonnier, 1955:172, pl. 48, figs. a–j; A. M. Clark and Rowe, 1971:185–186, pl. 30, fig. 8; Rowe and Doty, 1977:235– 236, figs. 5 c, 8 h; Cherbonnier, 1988:249– 251, fig. 111 a–j; Kerr, 1994:171; Rowe and Gates, 1995:338; Rowe and Richmond, 2004:3307–3308. material examined : uf 11413, Gilfith, < 1 m depth at night on rubble on middle reef flat, 14 December 2009. remarks : This species is a new record for Yap. The few other specimens of Euapta seen in this survey also all appear to be this species, rather than the closely related E . tahitiensis Cherbonnier, 1955, which also occurs in Mi- cronesia (Michonneau et al. 2013). This spe- cies was active at night on rubble and sand. Specimens seen during the day were hidden under rocks and rubble. Genus Opheodesoma Fisher, 1907 Opheodesoma grisea (Semper, 1868) Plate III F, G, H; Figure 5 D, E, F Synapta grisea Semper, 1868:11, pl. 4, figs. 6, 7. Opheodesoma grisea: Fisher, 1907:723; Heding, 1928:129, figs. 4.7, 7.3, 9; Cherbonnier, 1955:171, pl. 49, figs. k–t; A. M. Clark and Rowe, 1971:186, pl. 30, fig. 11; Rowe and Doty, 1977:235, figs. 5 a, 8 g; Cherbonnier, 1988:244 –246, fig. 109 a–f; Rowe and Gates, 1995:340; Rowe and Richmond, 2004:3308. material examined : uf 5862, 5876, Fanif, 10 m in blue hole, 3 August 2007. uf 5885, 5886, O’Keefe’s Island, 1 m depth in sea grass near massive Porites sp. colonies, 30 July 2007. uf 5855, 5900, O’Keefe’s Island, 1 m depth in sea grass near mangroves, 5 August 2007. uf 11406, Kaday, < 1 m depth during the day in sea grass bed on inner reef flat, 11 December 2009. remarks : This species is a new record for Yap. We found diverse color patterns among specimens referable to O. grisea (Plate III F, G, H ). This species was frequently observed in sea grass on inner reef flats, as well as at night in areas with high coral cover in the blue holes. Genus Polyplectana H. L. Clark, 1907 Polyplectana sp. Figure 5 G, H material examined : uf 5874, Maap, 2 m depth during the day in sea grass bed on inner reef flat, 29 July 2007. remarks : This species is a new record for Yap. This is a small species found in large numbers in sea grass, bordering a patch reef of Acropora spp. Absence of miliary granules and branched ossicles suggests that this spe- cies is P. kerfersteini. Genus Synapta Eschscholtz, 1829 Synapta maculata (Chamisso & Eysenhardt, 1821) Plate III E Holothuria maculata Chamisso & Eysenhardt, 1821:352, pl. 25; H. L. Clark, 1907:78, pls. 1, 4, figs. 17–19, 26; H. L. Clark, 1921:160; A. M. Clark, 1952:204; Cherbonnier, 1955:170, pl. 47, figs. a–d; A. M. Clark and Rowe 1971:186 –187, pl. 30, fig. 9. Synapta maculata andreae: Heding, 1928:115, text fig. 3, fig. 2. Synapta maculata sundaensis: Heding, 1928:116, text fig. 3, figs. 3–5. Synapta maculate: Heding, 1928:113, fig. 2; Rowe and Doty, 1977:234 –235, figs. 5 a, 8 e; Cherbonnier, 1988:107–108, fig. 43 a– k; Kerr, 1994:171; Rowe and Gates, 1995:343–344; Rowe and Richmond, 2004:3308. material examined : Photo record (Plate III E ), Gagil, 1.5 m depth in grass flats mixed with corals, 26 November 2002. uf 5894, Balabat, 15 m on sandy bottom of channel, 2 August 2007. uf 11424, Kaday, < 1 m depth during the day in sea grass bed on inner reef flat, 10 December 2009. remarks : This species is a new record for Yap. This large, diurnally active species was commonly found in sea grass. Specimens were also found rarely in silty sites, scattered with occasional coral colonies. Despite the rich fauna, only a handful of accounts report the holothuroids of Micro- nesia. The most recent and comprehensive account from Guam (Michonneau et al. 2013) enumerated 64 species of sea cucumbers, in- cluding 18 new records. The 37 species re- corded in this study is comparable to the di- versity known from Guam before the recent update (46 species) and makes Yap the best- documented Micronesian island after Guam. The known species composition of Yap is similar to that of Guam; the aspidochirotes dominate the reefs and adjacent habitats in Yap (86%) and Guam (81%), and the dendro- chirotes are the most poorly reported group in Yap (0%) and Guam (5%). Of the 37 holothuroid species recorded in this study, 24 (19 aspidochirotes and five apodans) are new records. At least two spe- cies, Actinopyga sp. and Holothuria ( Theelothuria ) sp., appear to be undescribed. One of the species we collected, H. ( Semperothuria ) roseomaculata , was also undescribed at the time of the surveys but was recently described (Kerr 2013). A few of the species were re ported only as photo records because we lacked permis- sion to collect during one visit. Two of these species, H . aff. ( Lessonothuria ) cavans and H . (? Thymiosycia ) sp., may represent rare species; however, we cannot make confident identifi- cations because we lack voucher specimens. The small-scale distribution patterns of holothuroids are closely related to micro- habitat and reef zonation (Levin 1979, Sloan 1979, 1982, Kerr et al. 1993, Kerr 1994). The distribution of Yapese holothuroids also varied considerably between reef zonations (Table 1). Sea grass beds on the inner reef flats were inhabited by the largest diversity of apodans, Opheodesoma grisea, Polyplectana sp., and Synapta maculata, as well as small cryptic aspidochirotes H . aff . ( Lessonothuria ) cavans and H . (? Thymiosycia ) sp., and the commer- cially valuable aspidochirotes Holothuria ( Metriatyla ) ? lessoni and H . ( Metriatyla ) scabra . The diurnally burying Bohadschia species, B. marmorata, B. vitiensis, and B. koellikeri, were also frequently observed in sea grass beds and near mangroves during the late afternoon and at night. Diurnally cryptic species, H. coluber, H. hilla, H. impatiens, H. pervicax, H. turriscelsa, Chiridota sp., Euapta godeffroyi, and Stichopus cf. monotuberculatus, were recorded on middle reef flats, where numerous coral colonies and rubble covered the bottom. Blue holes hosted an interesting set of less- common species, such as H . ( Cystipus ) inhabilis, H . ( Theelothuria ) sp., and H . ( Platyperona ) excellens . The most common species en- countered in the surveys, H. atra, was found at nearly all sites on inner and middle reef flats, and its large biomass was unrivaled by other species. Another common species, B. argus, was widespread at the western sites but was found less frequently at the more wave- exposed eastern sites. On the other hand, species that withstand higher wave energies, such as H. whitmaei and A. mauritiana, were found mostly on outer reef flats and edges at the eastern sites. In particular, A. mauritiana was absent at the relatively calm western sites. Table 1 summarizes the distribution pattern of Yapese holothuroids. The O’Keefe’s Island and Fanif sites were unique in having large sea grass beds and lack- ing middle and outer reef flats. The silty bot- tom of both sites was inhabited by Actinopyga sp. ...
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... ) chloronotos Brandt, 1835:50. Stichopus hirotai Mitsukuri, 1912:161, fig. 28. Stichopus chloronotus: Lampert, 1885:107; Théel, 1886:159, pl. 7, fig. 6; Sluiter, 1887:196; Ludwig, 1888:812; H. L. Clark, 1946:417; A. M. Clark and Rowe, 1971:178–179, pl. 27, fig. 18; Grosen- baugh, 1981:51–53; Cherbonnier, 1988:146 –147, fig. 60 a–o; Kerr, 1994:170; Rowe and Gates, 1995:377–378; Rowe and Richmond, 2004:3304. material examined : uf 5867, Colonia, 1 m depth on sand and rubble, 31 July 2007. uf 11407, Kaday, < 1 m depth during the day on rubble on middle reef flat, 11 December 2009. remarks : This species was common in almost all types of habitats from sea grass to inner and middle reef flats with high coral cover. It was also encountered on reef slopes at deeper depths, where individuals were often larger. Stichopus cf. monotuberculatus (Quoy & Gaimard, 1833) Figure 4 O, P, Q Holothuria monotuberculata Quoy & Gaimard, 1833:131, pl. 432, fig. I . Stichopus monotuberculatus: Cherbonnier, 1952:23, pl. 3, fig. 4, text fig. 8 a–t; Rowe and Gates, 1995:325; Samyn, 2000:15. Stichopus cf. monotuberculatus: Samyn and Vanden Berghe, 2000:31, pl. 2 F–H . material examined : uf 11415, Kaday, < 1 m depth during the day under rubble on middle reef flat, 11 December 2009. remarks : This species is a new record for Yap. This nocturnal species displayed an ar- ray of color patterns, also seen in other loca- tions in the Indo-Pacific (Byrne et al. 2010, Michonneau et al. 2013). The background varies from beige to camel with green and dark brown mottling. This species was com- monly observed under rocks and coral heads during the day and exposed on sand or coral- line rock at night. Recent genetic work by Byrne et al. (2010) revealed that S. horrens and S. monotuberculatus share similar sets of vari- able color patterns. The two species are dis- tinguished by ossicle morphology: S. horrens possesses tacklike body wall ossicles, whereas S. monotuberculatus does not (Byrne et al. 2010). The collected specimen was devoid of tacklike ossicles (Figure 4 O, P, Q ); however, ubiquity of the shared color forms in Yap sug- gests that S. horrens may also inhabit Yapese reefs. Stichopus vastus Sluiter, 1887 Stichopus vastus Sluiter, 1887:198–199, pl. 2, figs. 46 – 48; Sluiter, 1895:79; Rowe and Gates, 1995:381–382; Forbes et al., 1999:14; Massin, 1999:71–77, figs. 57 a–l, 58 a–m, 59 a–g, 60 a–d, 61, 112 d–e . Stichopus sp.: Allen and Steene, 1996:246; Gosliner et al., 1996:282, fig. 1039. material examined : uf 5367, 5869, 5913, O’Keefe’s Island, < 1 m depth during the day in sea grass bed near patchy reef, 30 July 2007. remarks : This species is a new record for Yap. It was uncommon and seen during the day and night on silty sand near mangroves and in the inner harbor. Genus Thelenota H. L. Clark, 1921 Thelenota ananas ( Jaeger, 1833) Trepang ananas Jaeger, 1833:24, pl. 3 fig. 1; H. L. Clark, 1922:48. Holothuria ananas: Quoy & Gaimard, 1833:110, pl. 6, figs. 1–3. Holothuria ( Thelenota ) ananas: Brandt, 1835:53. Stichopus ananas: Semper, 1868:75; Ludwig, 1882:133; Sluiter, 1901:30, pl. 2, fig. 1; Mitsukuri, 1912:150 –155, pl. 1, figs. 6 –8, text fig. 25. Thelenota ananas: A. M. Clark and Rowe, 1971:178–179, fig. 87 a; Rowe and Doty, 1977:227, figs. 2 b, 5 e; Grosenbaugh, 1981:51–53; Cherbonnier, 1988:152–153, fig. 63 a–k; Rowe and Gates, 1995:382. material examined : uf 5857, Balabat, 5 m depth during the day on pavement, 2 August 2007. remarks : This survey suggests that T. ananas is uncommon in Yap, relative to other locations in the western Pacific. It was oc- casionally seen exposed during the day and night on the outer reef slope on coralline rock or sand to at least 20 m deep. Thelenota rubralineata Massin & Lane, 1991 Plate III C Thelenota rubralineata Massin & Lane, 1991:57– 64, figs. 1–8; Kerr et al. 2007:21– 22, fig. 4. material examined : Photo record (Plate III C ), Ulithi Atoll, 25 m depth. Picture from Vanessa Fread, 27 July 2007. remarks : This species is a new record for Yap. Several specimens of this unmistakable and striking species have been seen by Brian Greene ( University of Hawai‘i at Mänoa, Ho- nolulu, pers. comm., 2007) around Yap and other islands in Yap State, on the deep outer reef slopes to at least 60 m depth. Order Apodida Family Chiridotidae Genus Chiridota Eschscholtz, 1829 Chiridota sp. Plate III D material examined : Photo record (Plate III D ), site unrecorded, exposed near coral rubble on middle reef flat, 28 November 2002. remarks : This species is a new record for Yap. This species usually takes shelter under rubble, often near sandy areas. Chiridota sp. often cleaves its body transversely when disturbed, as shown in Plate III D . This is likely either C . rigida Semper, 1868, or C. hawaiiensis Fisher, 1907, two widespread and poorly characterized species in the Pacific (A. M. Clark and Rowe 1971, Paulay 2003). Family Synaptidae Genus Euapta Oestergren, 1898 Euapta godeffroyi (Semper, 1868) Figure 5 A, B, C Synapta godeffroyi Semper, 1868:231, pl. 39, fig. 13. Euapta godeffroyi: Oestergren, 1898:113; H. L. Clark, 1907:72; H. L. Clark, 1921:158; Cherbonnier, 1955:172, pl. 48, figs. a–j; A. M. Clark and Rowe, 1971:185–186, pl. 30, fig. 8; Rowe and Doty, 1977:235– 236, figs. 5 c, 8 h; Cherbonnier, 1988:249– 251, fig. 111 a–j; Kerr, 1994:171; Rowe and Gates, 1995:338; Rowe and Richmond, 2004:3307–3308. material examined : uf 11413, Gilfith, < 1 m depth at night on rubble on middle reef flat, 14 December 2009. remarks : This species is a new record for Yap. The few other specimens of Euapta seen in this survey also all appear to be this species, rather than the closely related E . tahitiensis Cherbonnier, 1955, which also occurs in Mi- cronesia (Michonneau et al. 2013). This spe- cies was active at night on rubble and sand. Specimens seen during the day were hidden under rocks and rubble. Genus Opheodesoma Fisher, 1907 Opheodesoma grisea (Semper, 1868) Plate III F, G, H; Figure 5 D, E, F Synapta grisea Semper, 1868:11, pl. 4, figs. 6, 7. Opheodesoma grisea: Fisher, 1907:723; Heding, 1928:129, figs. 4.7, 7.3, 9; Cherbonnier, 1955:171, pl. 49, figs. k–t; A. M. Clark and Rowe, 1971:186, pl. 30, fig. 11; Rowe and Doty, 1977:235, figs. 5 a, 8 g; Cherbonnier, 1988:244 –246, fig. 109 a–f; Rowe and Gates, 1995:340; Rowe and Richmond, 2004:3308. material examined : uf 5862, 5876, Fanif, 10 m in blue hole, 3 August 2007. uf 5885, 5886, O’Keefe’s Island, 1 m depth in sea grass near massive Porites sp. colonies, 30 July 2007. uf 5855, 5900, O’Keefe’s Island, 1 m depth in sea grass near mangroves, 5 August 2007. uf 11406, Kaday, < 1 m depth during the day in sea grass bed on inner reef flat, 11 December 2009. remarks : This species is a new record for Yap. We found diverse color patterns among specimens referable to O. grisea (Plate III F, G, H ). This species was frequently observed in sea grass on inner reef flats, as well as at night in areas with high coral cover in the blue holes. Genus Polyplectana H. L. Clark, 1907 Polyplectana sp. Figure 5 G, H material examined : uf 5874, Maap, 2 m depth during the day in sea grass bed on inner reef flat, 29 July 2007. remarks : This species is a new record for Yap. This is a small species found in large numbers in sea grass, bordering a patch reef of Acropora spp. Absence of miliary granules and branched ossicles suggests that this spe- cies is P. kerfersteini. Genus Synapta Eschscholtz, 1829 Synapta maculata (Chamisso & Eysenhardt, 1821) Plate III E Holothuria maculata Chamisso & Eysenhardt, 1821:352, pl. 25; H. L. Clark, 1907:78, pls. 1, 4, figs. 17–19, 26; H. L. Clark, 1921:160; A. M. Clark, 1952:204; Cherbonnier, 1955:170, pl. 47, figs. a–d; A. M. Clark and Rowe 1971:186 –187, pl. 30, fig. 9. Synapta maculata andreae: Heding, 1928:115, text fig. 3, fig. 2. Synapta maculata sundaensis: Heding, 1928:116, text fig. 3, figs. 3–5. Synapta maculate: Heding, 1928:113, fig. 2; Rowe and Doty, 1977:234 –235, figs. 5 a, 8 e; Cherbonnier, 1988:107–108, fig. 43 a– k; Kerr, 1994:171; Rowe and Gates, 1995:343–344; Rowe and Richmond, 2004:3308. material examined : Photo record (Plate III E ), Gagil, 1.5 m depth in grass flats mixed with corals, 26 November 2002. uf 5894, Balabat, 15 m on sandy bottom of channel, 2 August 2007. uf 11424, Kaday, < 1 m depth during the day in sea grass bed on inner reef flat, 10 December 2009. remarks : This species is a new record for Yap. This large, diurnally active species was commonly found in sea grass. Specimens were also found rarely in silty sites, scattered with occasional coral colonies. Despite the rich fauna, only a handful of accounts report the holothuroids of Micro- nesia. The most recent and comprehensive account from Guam (Michonneau et al. 2013) enumerated 64 species of sea cucumbers, in- cluding 18 new records. The 37 species re- corded in this study is comparable to the di- versity known from Guam before the recent update (46 species) and makes Yap the best- documented Micronesian island after Guam. The known species composition of Yap is similar to that of Guam; the aspidochirotes dominate the reefs and adjacent habitats in Yap (86%) and Guam (81%), and the dendro- chirotes are the most poorly reported group in Yap (0%) and Guam (5%). Of the 37 holothuroid species recorded in this study, 24 (19 aspidochirotes and five apodans) are new records. At least two spe- cies, Actinopyga sp. and Holothuria ( Theelothuria ) sp., appear to be undescribed. One of the species we collected, H. ( Semperothuria ) roseomaculata , was also undescribed at the time of the surveys but was recently described (Kerr 2013). A few of the species were re ported only as photo records because we lacked permis- sion to collect during one visit. Two of these species, H . aff. ( Lessonothuria ) cavans and H . (? Thymiosycia ) sp., may represent rare species; however, we cannot make confident identifi- cations because we lack voucher specimens. The ...
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... Rowe and Richmond, 2004:3301. material examined : Photo record (Plate II A ), Gagil, < 1 m depth under exposed coral rubble on the outer reef flat at low tide, 28 November 2002. remarks : Although often abundant on other islands of Micronesia (e.g., Kerr et al. 1993, Kerr 1994), this species was seen in Yap only as a few small individuals under exposed coral rubble on the outer reef flat at low tide at Gagil. Holothuria ( Metriatyla ) ? lessoni Massin, Uthicke, Purcell, Rowe, & Samyn, 2009 Figure 3 G, H; Plate II B Holothuria timama Lesson, 1830:118, pl. 43; Lampert, 1885:94; Théel, 1886:240; A. M. Clark, 1963:383; Opinion 762, 1966:15; Melville and Smith, 1987:301. Holothuria ( Metriatyla ) timana: Rowe and Gates, 1995:295; Marsh and Morrison, 2004:339. Holothuria ( Metriatyla ) scabra: VandenSpiegel et al., 1992:168, figs. 2, 3 A–E, 4 A–G , non H. ( M. ) scabra Jaeger, 1833. Holothuria scabra var. versicolor Conand, 1986:19; Conand, 1991:170; Conand and Byrne, 1993:3ss; Forbes et al., 1999:38; Hamel et al., 2001:146, fig. 4 B; Uthicke et al., 2005:261ss, fig. 1 B–D . Holothuria aculeate: Cherbonnier, 1951:298, non H. aculeata Semper, 1868; Catala, 1979:245, fig. 91, non H. aculeata Semper, 1868; Rowe and Gates, 1995:295 (cited as a synonym of H. timana ). Holothuria ( Metriatyla ) aculeate: Rowe, 1969:160; A. M. Clark and Rowe, 1971:176. Holothuria ( Metriatyla ) lessoni Massin, Uthicke, Purcell, Rowe, & Samyn, 2009:41– 47, figs. 1, 3, 4, 5, table 1, 2. material examined : uf 5880, Gagil, < 1 m depth during the day in mangrove habitat near corals and sea grass, 6 August 2007. remarks : This species is a new record for Yap. One specimen was found in shallow water at the boundary between mangrove forest and sea grass bed with small, scattered Porites sp. The color pattern was gray with several small, black blotches (Plate II B ). This species has been frequently misidentified as H. scabra (see following species); however, the recent description of H. lessoni (Massin et al. 2009) clarified the diagnostic morpho- logical characters, such as size, color, and ossicles, to eliminate frequent confusion be- tween the two species. When describing, the authors were not able to use the name H. timama because it has been suppressed by ICZN Opinion 762 (1966). We are un- able to confidently iden tify the collected specimen because it shows mixed characters of H. lessoni and H. scabra. The collected spec- imen shows the diagnostic morphology of H. scabra including transverse grooves, lack of blotches, and general color pattern; however, the specimen also shows the characteristic, spiny table ossicles of H. lessoni (Figure 3 G, Plate II B ). Holothuria ( Metriatyla ) scabra Jaeger, 1833 Figure 3 I, J Holothuria scabra Jaeger, 1833:23. Holothuria ( Holothuria ) scabra: Panning, 1934 b: 80, fig. 66 a–f. Holothuria ( Metriatyla ) scabra: Rowe, 1969:160 –161, fig. 20 a–c; A. M. Clark and Rowe, 1971:178–179, fig. 871, pl. 15, fig. 15; Cherbonnier, 1980:647, fig. 16 A–L; Liao, 1980:116; Tan Tiu, 1981:83, pl. 25, figs. 1–3; Féral and Cherbonnier, 1986:86 – 87; Cherbonnier, 1988:135–137, fig. 55 a– o; Kerr, 1994:168, fig. 4 c; Rowe and Gates, 1995:294. material examined : uf 5851, O’Keefe’s Island, < 1 m depth during the day in sea grass at the edge of mangroves, 5 August 2007. remarks : This species is a new record for Yap. Several specimens were observed during this survey at a depth of about 1 m during the daytime. They all occurred on the inner reef flat, adjacent to mangroves, and in sea grass and on silty sand. Holothuria ( Microthele ) fuscogilva Cherbon- nier, 1980 Figure 3 K, L; Plate II C Holothuria ( Microthele ) fuscogilva Cherbon- nier, 1980:628, fig. 7, pl. C; Féral and Cherbonnier, 1986:88–89; Kerr et al., 1992:208, fig. 4 d, pl. 1 h . material examined : uf 11163, 11414, Kaday, < 1 m depth, one specimen collected during the day and one collected at night; both were in sea grass beds on the middle reef flat, 11 December 2009. remarks : This species is a new record for Yap. Holothuria fuscogilva is usually found at depths below 10 m; however, most of the specimens in Yap were observed at depths of less than 2 m in sea grass. Holothuria fuscogilva specimens in the western Pacific frequently display color patterns that are mixtures of black and white, hence the common name, white teatfish. Specimens seen in Yap showed a rather uniform beige background with dark brown mottling (Plate II C ). This color form has also been observed in other localities, such as Moorea (G. Paulay, pers. comm.), Guam (Kerr et al. 1992), Australia, and the Southwest Pacific ( Uthicke et al. 2004). Holothuria ( Microthele ) fuscopunctata Jaeger, 1833 Figure 3 M Holothuria fuscopunctata Jaeger, 1833:23; Sem- per, 1868:86, 277, pl. 30, fig. 29; Théel, 1886:235; Lampert, 1885:79; Ludwig, 1882:136. Holothuria axiologa H. L. Clark, 1921:175, pl. 38; Grosenbaugh, 1981:51. Holothuria ( Microthele ) fuscopunctata: Cher- bonnier, 1980:623, fig. 5; Rowe and Gates, 1995:361. material examined : uf 11390, Balabat, 3 m depth during the day on sand on middle reef flat, 14 December 2009. remarks : This large and distinctive spe- cies was observed occasionally on sandy mid- dle reef flats at depths between 1 and 3 m. Holothuria ( Microthele ) whitmaei Bell, 1887 Figure 3 N, O Holothuria ( Bohadschia ) whitmaei Bell, 1887:532–533, pl. 45, fig. 4. Holothuria mammifera Saville-Kent, 1890:4, pl. 1, fig. 3. Holothuria ( Microthele ) nobilis Selenka, 1867:313, pl. 17, fig. 16; Rowe, 1969:162, 164, fig. 21; A. M. Clark and Rowe, 1971:178–179, fig. 87 m, pl. 27, fig. 10, pl. 28, fig. 20; Rowe and Doty, 1977:231, figs. 3 f, 7 d; Grosenbaugh, 1981:51–53; Cherbonnier, 1988:142–144, fig. 58 a–l . Holothuria ( Microthele ) whitmaei: Rowe and Gates, 1995:362. material examined : uf 5866, Tamil, 1 m depth on rubble on middle reef flat, 31 July 2007. uf 11389, Gilfith, 3 m depth during the day on rubble on middle reef flat, 13 De- cember 2009. remarks : This species was commonly found on the eastern outer reef flats and the western reef edges. This species tends to in- habit high-energy habitats with strong wave action. Adults of this species resemble H. fus- cogilva in shape and size but have a uniformly black dorsum. Holothuria ( Platyperona ) excellens (Ludwig, 1875) Plate II D M lleria excellens Ludwig, 1875:98, fig. 32 a–c; Lampert, 1885:97. Holothuria ( Microthele ) excellens: Panning, 1929:132, fig. 16. Holothuria ( Platyperona ) excellens: Théel, 1886:199; Cherbonnier, 1988:94 –95, fig. 37 a–n; Kerr et al., 1992:209–213, pl. 1 e–f, figs. 3 g–h, 4 a–c, 5 a–e . material examined : uf 5879, Tamil blue hole, 15 m depth at night, 30 July 2007. remarks : This species is a new record for Yap. One specimen was seen at night in an area with high coral cover. This species is uniformly brown with a purplish tinge, has a smooth body wall, and ejects numerous fine Cuvierian tubules when handled (Plate II D ). Holothuria ( Semperothuria ) roseomaculata Kerr, 2013 Plate II E; Figure 4 A, B Holothuria flavomaculata: Yamanouti, 1939:604 – 634; Yamanouti, 1956:361; Amesbury et al., 1977:14 –16; Grosen- baugh, 1981:51–53; Colin and Arneson, 1995:260, fig. 1232. Holothuria ( Semperothuria ) flavomaculata: Féral and Cherbonnier, 1986:90 –91, fig. 40 D, unnumb. fig. (p. 90); Conand, 1989:27 et non seq., figs. 18–19; Purcell et al., 2013:50 –51, unnumb. figs. Holothuria ( Semperothuria ) sp.: Kerr et al., 2007:15, 31, fig. 3 b . Holothuria ( Semperothuria ) non flavomaculata: Friedman et al., 2008:8, 12; Tardy and Pakoa, 2009:10, 49–50. Holothuria ( Semperothuria ) roseomaculata Kerr, 2013:384 –394. material examined : uf 5849, 5850, 5872, 5890, 5911, O’Keefe’s Island, < 1 m depth in sea grass near patchy massive Porites sp., 30 July 2007. uf 11397, 11403, Fanif, < 1 m depth during the day under a massive Porites sp. on inner reef flat, 12 December 2009. remarks : This species was locally abun- dant near O’Keefe’s Island and Fanif in sea grass. This species has been misidentified as H. flavomaculata, of which the color and form are consistently different from those of this species. Holothuria flavomaculata possesses yellow papillae, whereas H . ( S. ) roseomaculata has larger, rose-colored papillae (Plate II E ). We found this species on silty sand, under massive Porites spp. colonies near O’Keefe’s Island, at Fanif, and in the Colonia harbor. Based on the similarity in abundance and hab- itat, it appears to be the “Holothuria moebii” mentioned in Amesbury et al. (1976). Holothuria ( Stauropora ) pervicax Selenka, 1867 Figure 4 C, D Holothuria pervicax Selenka, 1867:327, pl. 18, fig. 54; A. M. Clark, 1952:204. Holothuria depressa Ludwig, 1875:108, fig. 44. Holothuria ( Mertensiothuria ) pervicax: Rowe, 1969:149; A. M. Clark and Rowe, 1971:176 –177; Rowe and Doty, 1977:234, figs. 4 g, 8 c; A. M. Clark, 1984:99; Cher- bonnier, 1988:107–108, fig. 43 a–k; Kerr, 1994:168. Holothuria ( Stauropora ) pervicax: Rowe and Gates, 1995:368; Rowe and Richmond, 2004:3303. material examined : uf 11425, Kaday, < 1 m depth at night on sand under a coral head on middle reef flat, 11 December 2009. remarks : This species is a new record for Yap. Holothuria pervicax was an uncommon nocturnal species on Yap and seen only in Kaday. Specimens occurred under rocks and coral heads during the day and were seen ex- posed on sand and rubble at night. Holothuria ( Theelothuria ) sp. Plate II F, G; Figure 4 E, F material examined : uf 5892, Tamil blue hole, 15 m depth at night in blue hole among dead coral rubble, 31 July 2007. uf 5893, Gagil blue hole, 12 m depth at night in blue hole, 6 August 2007. uf 5896, Fanif blue hole, 15 m depth at night in blue hole, 3 August 2007. remarks : This species is a new record for Yap. It is a relatively common holothuroid at night in the blue holes on the western reefs (Plate II F, G ). Most specimens possessed a gray dorsum with many raised tubercles and a slightly ...
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... fig. 44. Holothuria ( Mertensiothuria ) pervicax: Rowe, 1969:149; A. M. Clark and Rowe, 1971:176 –177; Rowe and Doty, 1977:234, figs. 4 g, 8 c; A. M. Clark, 1984:99; Cher- bonnier, 1988:107–108, fig. 43 a–k; Kerr, 1994:168. Holothuria ( Stauropora ) pervicax: Rowe and Gates, 1995:368; Rowe and Richmond, 2004:3303. material examined : uf 11425, Kaday, < 1 m depth at night on sand under a coral head on middle reef flat, 11 December 2009. remarks : This species is a new record for Yap. Holothuria pervicax was an uncommon nocturnal species on Yap and seen only in Kaday. Specimens occurred under rocks and coral heads during the day and were seen ex- posed on sand and rubble at night. Holothuria ( Theelothuria ) sp. Plate II F, G; Figure 4 E, F material examined : uf 5892, Tamil blue hole, 15 m depth at night in blue hole among dead coral rubble, 31 July 2007. uf 5893, Gagil blue hole, 12 m depth at night in blue hole, 6 August 2007. uf 5896, Fanif blue hole, 15 m depth at night in blue hole, 3 August 2007. remarks : This species is a new record for Yap. It is a relatively common holothuroid at night in the blue holes on the western reefs (Plate II F, G ). Most specimens possessed a gray dorsum with many raised tubercles and a slightly lighter ventrum (Plate II F ). A less- common color morph also seen had an olive green dorsum (Plate II G ) but was otherwise identical in form, habitat, and behavior. Dorsal and ventral body wall ossicles consist of tables and buttons. Tables have flattened, lacelike disks with spiny margins and a large central hole in which the four tiers form a dis- tinctive cross. Buttons are variably developed and often knobby. When disturbed, speci- mens readily ejected many long translucent Cuvierian tubules, most resembling those of H . ( Theelothuria ) turriscelsa and H . ( Stauropora ) pervicax . Holothuria ( Theelothuria ) turriscelsa Cherbon- nier, 1980 Figure 4 G, H Holothuria ( Theelothuria ) turriscelsa Cher- bonnier, 1980:644; pl. 1 e, fig. 15 a–l; Féral and Cherbonnier, 1986:92–93; Kerr et al., 1992:209. material examined : uf 5878, Tamil, 15 m depth at outer reef slope, 2 August 2007. uf 11396, Kaday, < 1 m depth at night under a coral head on middle reef flat, 11 December 2009. remarks : This species is a new record for Yap. It was found under coral heads and rubble on middle reef flats in Kaday. At night, H. turriscelsa was often seen roaming on the reef flat. Specimens readily ejected thick, transparent Cuvierian tubules when disturbed. Holothuria ( Thymiosycia ) impatiens (Forsskål, 1775) Plate II H; Figure 4 I, J; Plate III A Fistularia impatiens Forsskål, 1775:121, pl. 39 b . Holothuria impatiens: Haacke, 1880:46; H. L. Clark, 1921:178, pl. 19, figs. 3, 5; Panning, 1935:86 –88, fig. 72; Panning, 1941:7, figs. 5– 6. Holothuria impatiens concolor H. L. Clark, 1921:179. Holothuria impatiens lutea H. L. Clark, 1921:179. Holothuria impatiens pulchra H. L. Clark, 1921:179, pl. 19. Holothuria impatiens bicolor H. L. Clark, 1938:522. Holothuria ( Thymiosycia ) impatiens: Rowe, 1969:146 –147, fig. 13 a–c; A. M. Clark and Rowe, 1971:178–179, fig. 85 a–a ′ , pl. 26, fig. 2, pl. 28, fig. 8; Rowe and Doty, 1977:233, figs. 4 c, 7 e; Cherbonnier, 1988:88–89, fig. 35 a–k; Kerr, 1994:170; Rowe and Gates, 1995:372–373; Rowe and Richmond, 2004:3303–3304. material examined : uf 5877, Kaday, blue hole, 4 August 2007. uf 11427, Kaday, < 1 m depth during the day on sand under a rock on middle reef flat, 11 December 2009. remarks : This species is a new record for Yap. This cryptic species was found in- frequently under rocks at the western sites Kaday and Gilfith. As for many other holo- thuroids, the taxonomic status of H. impatiens has been debated for years because of its color and morphological variation. It is likely a complex of several species (F. Michonneau, pers. comm.). We found at least three distinct color variants with morphology and ossicle variation currently assignable to H. impatiens (Plates II H, III A ). One of the forms was nearly all white and inhabited the blue holes of the reef flat. It was nocturnally active and retracted into its shelter at even the slightest exposure to light, making it difficult to photo- graph in situ. Hence, we do not have a photo record of this variant. Holothuria (? Thymiosycia ) sp. Plate III B material examined : Photo record (Plate III B ), site unrecorded, < 1 m depth in sea grass, collected by E. Tardy, 16 Septem- ber 2009. remarks : This species is a new record for Yap. The uncollected photographed specimen’s mustard color, sparsely placed pa- pillae, and habitat resemble those of a form of H . ( Thymiosycia ) impatiens that has also been observed in Palau (A. M. Kerr, unpubl. obs.). Genus Pearsonothuria Levin, Kalinin, & Stonik, 1984 Pearsonothuria graeffei (Semper, 1868) Figure 4 K, L Holothuria graeffei Semper, 1868:78, pl. 30, fig. 9 a–b . Bohadschia drachi Cherbonnier, 1955:134, pl. 24 a–n . Bohadschia graeffei: Panning, 1929:124, fig. 5; Panning, 1944:44, fig. 13; A. M. Clark and Rowe, 1971:176 –177, pl. 27, fig. 7; Rowe and Doty, 1977:229, figs. 2 g, 6 e . Pearsonothuria graeffei: Levin et al. 1984:33– 38, figs. 1, 2; Cherbonnier, 1988:49–50, fig. 17 a–f; Kerr, 1994:170; Rowe and Gates, 1995:376. material examined : uf 5895, Kaday, 15 m depth at outer reef slope, 2 August 2007. uf 11393, Colonia outer reef, 10 m depth on coral on reef slope, 15 December 2009. remarks : This species is a new record for Yap. In this survey, this species was found only at the Colonia outer reef slope around 10 m in depth, but it is a conspicuous and abundant species in other locations in the western Pacific (Massin 1999). Family Stichopodidae Genus Stichopus Brandt, 1835 Stichopus chloronotus Brandt, 1835 Figure 4 M, N Stichopus ( Perideris ) chloronotos Brandt, 1835:50. Stichopus hirotai Mitsukuri, 1912:161, fig. 28. Stichopus chloronotus: Lampert, 1885:107; Théel, 1886:159, pl. 7, fig. 6; Sluiter, 1887:196; Ludwig, 1888:812; H. L. Clark, 1946:417; A. M. Clark and Rowe, 1971:178–179, pl. 27, fig. 18; Grosen- baugh, 1981:51–53; Cherbonnier, 1988:146 –147, fig. 60 a–o; Kerr, 1994:170; Rowe and Gates, 1995:377–378; Rowe and Richmond, 2004:3304. material examined : uf 5867, Colonia, 1 m depth on sand and rubble, 31 July 2007. uf 11407, Kaday, < 1 m depth during the day on rubble on middle reef flat, 11 December 2009. remarks : This species was common in almost all types of habitats from sea grass to inner and middle reef flats with high coral cover. It was also encountered on reef slopes at deeper depths, where individuals were often larger. Stichopus cf. monotuberculatus (Quoy & Gaimard, 1833) Figure 4 O, P, Q Holothuria monotuberculata Quoy & Gaimard, 1833:131, pl. 432, fig. I . Stichopus monotuberculatus: Cherbonnier, 1952:23, pl. 3, fig. 4, text fig. 8 a–t; Rowe and Gates, 1995:325; Samyn, 2000:15. Stichopus cf. monotuberculatus: Samyn and Vanden Berghe, 2000:31, pl. 2 F–H . material examined : uf 11415, Kaday, < 1 m depth during the day under rubble on middle reef flat, 11 December 2009. remarks : This species is a new record for Yap. This nocturnal species displayed an ar- ray of color patterns, also seen in other loca- tions in the Indo-Pacific (Byrne et al. 2010, Michonneau et al. 2013). The background varies from beige to camel with green and dark brown mottling. This species was com- monly observed under rocks and coral heads during the day and exposed on sand or coral- line rock at night. Recent genetic work by Byrne et al. (2010) revealed that S. horrens and S. monotuberculatus share similar sets of vari- able color patterns. The two species are dis- tinguished by ossicle morphology: S. horrens possesses tacklike body wall ossicles, whereas S. monotuberculatus does not (Byrne et al. 2010). The collected specimen was devoid of tacklike ossicles (Figure 4 O, P, Q ); however, ubiquity of the shared color forms in Yap sug- gests that S. horrens may also inhabit Yapese reefs. Stichopus vastus Sluiter, 1887 Stichopus vastus Sluiter, 1887:198–199, pl. 2, figs. 46 – 48; Sluiter, 1895:79; Rowe and Gates, 1995:381–382; Forbes et al., 1999:14; Massin, 1999:71–77, figs. 57 a–l, 58 a–m, 59 a–g, 60 a–d, 61, 112 d–e . Stichopus sp.: Allen and Steene, 1996:246; Gosliner et al., 1996:282, fig. 1039. material examined : uf 5367, 5869, 5913, O’Keefe’s Island, < 1 m depth during the day in sea grass bed near patchy reef, 30 July 2007. remarks : This species is a new record for Yap. It was uncommon and seen during the day and night on silty sand near mangroves and in the inner harbor. Genus Thelenota H. L. Clark, 1921 Thelenota ananas ( Jaeger, 1833) Trepang ananas Jaeger, 1833:24, pl. 3 fig. 1; H. L. Clark, 1922:48. Holothuria ananas: Quoy & Gaimard, 1833:110, pl. 6, figs. 1–3. Holothuria ( Thelenota ) ananas: Brandt, 1835:53. Stichopus ananas: Semper, 1868:75; Ludwig, 1882:133; Sluiter, 1901:30, pl. 2, fig. 1; Mitsukuri, 1912:150 –155, pl. 1, figs. 6 –8, text fig. 25. Thelenota ananas: A. M. Clark and Rowe, 1971:178–179, fig. 87 a; Rowe and Doty, 1977:227, figs. 2 b, 5 e; Grosenbaugh, 1981:51–53; Cherbonnier, 1988:152–153, fig. 63 a–k; Rowe and Gates, 1995:382. material examined : uf 5857, Balabat, 5 m depth during the day on pavement, 2 August 2007. remarks : This survey suggests that T. ananas is uncommon in Yap, relative to other locations in the western Pacific. It was oc- casionally seen exposed during the day and night on the outer reef slope on coralline rock or sand to at least 20 m deep. Thelenota rubralineata Massin & Lane, 1991 Plate III C Thelenota rubralineata Massin & Lane, 1991:57– 64, figs. 1–8; Kerr et al. 2007:21– 22, fig. 4. material examined : Photo record (Plate III C ), Ulithi Atoll, 25 m depth. Picture from Vanessa Fread, 27 July 2007. remarks : This species is a new record for Yap. Several specimens of this unmistakable and striking species have been seen by Brian Greene ( University of Hawai‘i at Mänoa, Ho- nolulu, pers. comm., ...
Context 6
... ( Bohadschia ) argus: Panning, 1929:121, fig. 50 a–c . material examined : uf 5873, Maap, < 1 m depth during the day on sand on inner reef flat, 29 July 2007. uf 11417, Kaday, < 1 m depth during the day on sand on inner reef flat, 11 December 2009. remarks : This species was found at all but two sites located between the volcanic is- lands, Fanif and O’Keefe’s Island, where the bottom consisted of soft mud and sea grass. The most common color pattern observed in Yap was a gray background with dark ocella- tions, and others were dark brown or, rarely, pure white with only faint gray ocellations. Bohadschia marmorata Jaeger, 1833 Figure 2 G, H Bohadschia marmorata Jaeger, 1833:18, pl. 3, fig. 9; Brandt, 1835:56; A. M. Clark and Rowe, 1971:176 –177, 209, pl. 27, fig. 8; Rowe and Doty, 1977:229–230, figs. 3 a, 6 g, h; Cherbonnier, 1988:36, fig. 11 a – l; Kim et al., 2013:88–89, fig. 3 K . Sporadipus ualanensis Brandt, 1835:46. Holothuria ualensis Selenka, 1867:341. Holothuria marmorata: Semper, 1868:79, pl. 80, fig. 10, pl. 35, fig. 3, pl. 36, fig. 8, pl. 37, figs. 1– 4; Lampert, 1885:36, 86 –87; Théel, 1886:202–203; H. L. Clark, 1938:523. Holothuria ( Bohadschia ) marmorata: Panning, 1929:120, fig. 1. material examined : uf 5856, 5859, 11401, Kaday, < 1 m depth at night in sea grass near mangroves, and on sand on middle reef flat, 5– 6 August 2007, 11 December 2009. remarks : This species is a new record for Yap. This species was most frequently observed at night in sea grass and sand flats of the inner and middle reef flats. Bohadschia marmorata has frequently been referred to other species within the genus in the litera- ture. However, a recent study by Kim et al. (2013) revealed that B. marmorata sensu Jaeger is genetically distinct from other spe- cies frequently misidentified as B. marmorata . In addition, B. marmorata can be easily distin- guished by its dorsolateral markings and rela- tively small size (Kim et al. 2013). Bohadschia koellikeri (Semper, 1868) Figure 2 I, J; Plate I D, F Holothuria koellikeri Semper, 1868:86, pl. 30, fig. 25, pl. 35, fig. 7; Lampert, 1885:36, 86 –87; Théel, 1886:204; Rowe and Gates, 1995:351. Bohadschia koellikeri: Rowe, 1969:130; A. M. Clark and Rowe, 1971:194, 209; Kim et al., 2013:86 –88, figs. 3 E, F . material examined : uf 11420, Kaday, < 1 m depth at night on sand on inner reef flat, 10 December 2009. remarks : This species was nocturnal and only found in Kaday. Bohadschia koellikeri has been frequently misidentified as Bohadschia vitiensis, a morphologically similar but ge- netically distinct species (Kim 2010, Kim et al. 2013). Fine lineation and light brown blotches on the dorsal body wall clearly sepa- rate B. koellikeri from B. vitiensis, which pos- sesses a smooth dorsum, often with solid transverse bands (see Plate I D, E, F, G ). The overall color pattern of B. koellikeri observed in Yap was creamy beige with light brown blotches on the dorsum (Plate I D ). Bohadschia vitiensis (Semper, 1868) Figure 2 K, L; Plate I E, G Holothuria vitiensis Semper, 1868:80, pl. 30, fig. 12; Lampert, 1885:89; Théel, 1886:203–204; Pearson, 1913:57– 60, pl. VII, fig. 6; Domantay, 1933:76, pl. 1, fig. 2; Domantay, 1953:119. Holothuria similis Semper, 1868:85–86, pl. 25, 30. Holohuria clemens Ludwig, 1875:107, fig. 49. Holothuria bivittata Mitsukuri, 1912:68–71, pl. 3, fig. 75. Bohadschia marmorata vitiensis: Panning, 1944:40, fig. 11 a–y . Bohadschia vitiensis: Pearson, 1914:170; Rowe, 1969:130; A. M. Clark and Rowe, 1971:194; Rowe and Doty, 1977:229–230; Cher- bonnier, 1988:42– 44, fig. 14 a–i ; Kerr, 1994:167–168; Rowe and Richmond, 2004:3300; Kim et al., 2013:89, fig. 3 G–J . material examined : uf 5852, 5853, 5914, Colonia, 1 m depth in sea grass at edge of mangroves, 5 August 2007. uf 5891, 5915, Tamil, 2 m depth on sand on middle reef flat, 31 July 2007. uf 11394, Kaday, < 1 m depth at night on sand on inner reef flat, 10 December 2009. remarks : This species was found in Kaday and Tamil at night. The specimens were buried in sand during the day and emerged during the late afternoon. They were mostly found on inner reef flats in sea grass or sand flats in large numbers. Bohadschia vitiensis showed a wide variation in shad- ing. The background coloration varied from creamy white to brown, and the two dorsal transverse bands could be absent or quite distinct (Plate I E ). Bohadschia vitiensis has fre- quently been misidentified as B. marmorata; however, a study by Clouse et al. (2005) re- vealed that specimens referable to B. bivittata Mitsukuri, 1912, were genetically distinct and deserved species status. A recent study by Kim et al. (2013) added that color forms matching the description of B. bivittata fell within the B. vitiensis clade, in which the members show variable color and banding patterns. Genus Holothuria Linnaeus, 1767 Holothuria ( Acanthotrapezia ) coluber Semper, 1868 Figure 2 M Holothuria coluber Semper, 1868:90, pl. 28, pl. 30, fig. 28 a, b, pl. 34, fig. 5; Panning, 1944:62, fig. 30 a–i . Holothuria ( Holothuria ) coluber: Panning, 1934 a :35, fig. 30 a, b . Holothuria ( Acanthotrapezia ) coluber: Rowe, 1969:138–139; A. M. Clark and Rowe 1971:176 –177, fig. 85 c, pl. 27, fig. 13; Levin, 1979:20; Cherbonnier, 1980:636, fig. 11 A–H; Tan Tiu, 1981:74, pl. 16, figs. 1–3; Cannon and Silver, 1986:21, fig. 6 c ; Féral and Cherbonnier, 1986:80 –81, fig. 40 f ; Kerr, 1994:168, fig. 4; Rowe and Gates, 1995:290. material examined : uf 5888, 11419, O’Keefe’s Island, < 1 m depth during the day under a massive Porites sp. on inner reef flat, 30 July 2007, 13 December 2009. remarks : This species was found only near O’Keefe’s Island. During the day, the observed specimens were less active, often hiding under rocks or coral heads. The speci- mens extended their anterior end to feed at night. This may be the “Holothuria leucospi- lota” of Amesbury et al. (1977) and Grosen- baugh (1981), who noted a species in identical habitat also extending its anterior end to feed, especially in the evening. Holothuria ( Cystipus ) inhabilis Selenka, 1867 Figure 2 N, O; Plate I H Holothuria inhabilis Selenka, 1867:333, pl. 19, figs. 73–74. Jaegerothuria inhabilis: Deichmann, 1958:323– 324, pl. 8, figs. 14 ...

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