Microporella tonkinensis Dick & Ngai & Doan 2020, n. sp.

Microporella tonkinensis n. sp. (Fig. 12 A–D) urn:lsid:zoobank.org:act: D8480B51-CC0D-4B7B-B038-581C87933ADC Etymology. The specific name comes from the Gulf of Tonkin, wherein lies the Co To island group. Material examined. Holotype, VNMN-0248 (CT-6); paratype, VNMN-0249 (CT-41); both on SEM stubs....

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Main Authors: Dick, Matthew H., Ngai, Nguyen Danh, Doan, Hung Dinh
Format: Text
Language:unknown
Published: Zenodo 2020
Subjects:
Online Access:https://dx.doi.org/10.5281/zenodo.3703680
https://zenodo.org/record/3703680
id ftdatacite:10.5281/zenodo.3703680
record_format openpolar
institution Open Polar
collection DataCite Metadata Store (German National Library of Science and Technology)
op_collection_id ftdatacite
language unknown
topic Biodiversity
Taxonomy
Animalia
Bryozoa
Gymnolaemata
Cheilostomatida
Microporellidae
Microporella
Microporella tonkinensis
spellingShingle Biodiversity
Taxonomy
Animalia
Bryozoa
Gymnolaemata
Cheilostomatida
Microporellidae
Microporella
Microporella tonkinensis
Dick, Matthew H.
Ngai, Nguyen Danh
Doan, Hung Dinh
Microporella tonkinensis Dick & Ngai & Doan 2020, n. sp.
topic_facet Biodiversity
Taxonomy
Animalia
Bryozoa
Gymnolaemata
Cheilostomatida
Microporellidae
Microporella
Microporella tonkinensis
description Microporella tonkinensis n. sp. (Fig. 12 A–D) urn:lsid:zoobank.org:act: D8480B51-CC0D-4B7B-B038-581C87933ADC Etymology. The specific name comes from the Gulf of Tonkin, wherein lies the Co To island group. Material examined. Holotype, VNMN-0248 (CT-6); paratype, VNMN-0249 (CT-41); both on SEM stubs. Measurements. Holotype , VNMN-0248: AzL , 0.43–0.59 (0.513 ± 0.050); AzW , 0.28–0.48 (0.357 ± 0.051); OrL , 0.062 –0.074 (0.068 ± 0.003); OrW , 0.106 –0.124 (0.113 ± 0.005); OvL , 0.19–0.23 (0.207 ± 0.011); OvW , 0.23–0.27 (0.260 ± 0.011); AvRL , 0.090 –0.114 (0.102 ± 0.007); AvRW , 0.046 –0.063 (0.053 ± 0.006) (n = 15, 1). Paratype , VNMN-0249: AzL , 0.34–0.49 (0.428 ± 0.039); AzW , 0.23–0.38 (0.294 ± 0.043); OrL , 0.056 –0.072 (0.066 ± 0.004); OrW , 0.087 –0.104 (0.098 ± 0.005); AvRL , 0.068 –0.092 (0.082 ± 0.007); AvRW , 0.038 –0.057 (0.046 ± 0.005) (n = 15, 1). Diagnosis. Orifice with straight, weakly denticulate proximal margin; small condyles mediad to proximolateral corners of orifice. Modal number of oral spines four or five; spine bases often evident in zooids away from colony margin. Ascopore non-cribriform, without conspicuous umbo; pustulation between ascopore and orifice. Avicularia lateral or proximolateral to ascopore, one per zooid, small relative to orifice size. Non-personate, non-ribbed ooecium retaining pair of spines at proximolateral corners. Description. Colony unilaminar, encrusting, sheet-like. Zooids irregularly hexagonal, distinct, delineated by groove. Frontal wall markedly convex, with two to five small, inconspicuous areolae along each lateral margin; perforated over entire surface by small pseudopores, except in zone between ascopore and orifice; texture uniformly coarsely pustulose between pores; pustules usually present between ascopore and orifice, sometimes in discrete row. Orifice broader than long, broadly D-shaped, surrounded by low, smooth rim; proximal margin straight or slightly concave between conspicuous triangular condyles; condyles positioned slightly mediad to proximolateral corners of orifice; proximal margin weakly denticulate. Oral spines short, articulated; three (n = 5) or four (n = 27) in colony CT-6; three (n = 1), four (n = 13), five (n = 17), or six (n = 4) in colony CT-41; spines retained even in older parts of colony. Ascopore small, broader than long, encircled by smooth, raised rim, separated from proximal orificial margin by 1–1.5 times height of ascopore plus rim; with small lingula projecting proximally, leaving C-shaped opening; margin of opening finely denticulate all around. Sometimes proximal part of ascopore rim widened and raised as low, smooth tubercle. One avicularium per zooid, lateral to ascopore and directed distolaterally or sometimes laterally, or along zooidal margin proximolateral to ascopore, sometimes at lateral corner of zooid or even more proximally, directed distolaterally or laterally. Frontal avicularia moderately small, rostrum raised, sides and edges smooth, crossbar complete, area proximal to crossbar semicircular, mandibular area triangular, slightly longer than broad, guttered distally to allow for setiform mandible; setiform portion of mandible as long as or shorter than triangular rostral portion. Ooecium prominent, resting on frontal wall of next-distal zooid, broader than long, closed by operculum, with narrow, smooth, flat zone around proximal margin where operculum meets ooecium; with many small pseudopores around steep periphery, few in central-proximal area; top more finely pustulose than frontal wall, coarsely pustulose around periphery, with some moderately large marginal openings at base around distal margin; ooecium appearing peripherally ribbed at low magnification. Most-proximal pair of spines lateral to orifice retained in ooecium-bearing zooids, abutting proximolateral corners of ooecium. Remarks. Microporella is a speciose genus, with roughly 140 mostly Recent species described worldwide, from polar regions to the tropics (Bock 2018). The generic features are rather stereotyped, simplifying identification to genus but complicating identification to species. Until relatively recently, taxonomists viewed the European species M. ciliata (Pallas, 1766) as having a cosmopolitan distribution that essentially mirrored the range of the genus. While many species worldwide were correctly delineated from M. ciliata on the basis of unique apomorphies or combinations of characters, in the older literature many specimens from the tropical Pacific and elsewhere more or less similar to M. ciliata were attributed to that species, without description; for example, Canu & Bassler (1929) list M. ciliata from Hong Kong and the Philippines. Some of the characters utilized in the taxonomy of this genus are variable rather than discrete. For example, M. germana Dick & Ross, 1988 at Kodiak, Alaska, had three to five oral spines, with the modal number four, but at Ketchikan had four to six spines, with the modal number five (Dick et al . 2005). Similar interpopulation variation is seen in the frequencies of zooids having single, paired, or no avicularia per zooid, and in the shape of the ascopore (e.g., Harmelin et al . 2011, p. 18, fig. 9). One useful character that distinguishes one large group of species from another is the presence or absence of a personate ooecium (ooecium-bearing zooids have a raised peristomial rim, proximal and lateral to the orifice, that meets the ooecium distally) (Harmelin et al . 2011); M. dentilingua above has a personate ooecium, whereas M. tonkinensis n. sp. does not. Several similar species in the northwestern Pacific differ from M. tonkinensis n. sp. in the following ways. Microporella formosa Suwa & Mawatari, 1998 from Hokkaido, Japan, has larger avicularia relative to orifice size; the proximal orificial margin is smooth, with the condyles larger and occupying the proximolateral corners of the orifice. The orifice in Microporella antiborealis Liu & Liu, 2001 [listed in Liu et al . (2001) as previously described rather than as a new species, but we could find no reference for Liu & Liu (2001)] from China has a similarly finely denticulate proximal margin, but has larger avicularia relative to orifice size, a larger ascopore, and modal spine number six, and lacks the pair of lateral oral spines remaining at the proximolateral corners of the ooecium. Like M. tonkinensis n. sp. , Microporella borealis Suwa & Mawatari, 1998 from Hokkaido, Japan, has a finely denticulate proximal orificial margin, and has four or five oral spines, retaining one pair at the proximolateral corners of the ooecium, but the condyles are larger and in the proximolateral corners of the orifice, the avicularia are much larger relative to orifice size, and there is no pustulation between the ascopore and orifice. Only two specimens of M. tonkinensis were found; these differed markedly in zooid size, with CT-6 having larger zooids than CT-41. This sort of variation seems to be common in tropical to subtropical cheilostomes, both within and between colonies; a similar case was observed in Rhynchozoon lunifrons Dick & Grischenko, 2017 at Okinawa. While some of this variation may be due to differences in temperature at time of growth (O’Dea & Okamura 1999, 2000), the degree of variation seems greater than occurs in cold-temperate faunas, and intercolony variation may require additional explanation. In any case, bryozoan taxonomists working in the tropics and subtropics should be aware that zooid size should be treated with caution as a taxonomic character. Distribution. Co To Island is the only known locality. : Published as part of Dick, Matthew H., Ngai, Nguyen Danh & Doan, Hung Dinh, 2020, Taxonomy and diversity of coelobite bryozoans from drift coral cobbles on Co To Island, northern Vietnam, pp. 201-252 in Zootaxa 4747 (2) on pages 227-228, DOI: 10.11646/zootaxa.4747.2.1, http://zenodo.org/record/3694740 : {"references": ["Bock, P. (2018) Indexes to bryozoan taxa. Available from: http: // www. bryozoa. net / indexes. html (accessed 15 December 2018)", "Pallas, P. S. (1766) Elenchus Zoophytorum Sistens Generum Adumbrationes Generaliores et Speciarum Cognitarum Cuccintas Descriptiones cum Selectis Auctorus Synonymis. Petrum van Cleef, Hagae-Comitum, 451 pp. https: // doi. org / 10.5962 / bhl. title. 6595", "Canu, F. & Bassler, R. S. (1929) Bryozoa of the Philippine region. United States National Museum Bulletin, 100, I-XI + 1 - 685.", "Dick, M. H. & Ross, J. R. P. (1988) Intertidal Bryozoa (Cheilostomata) of the Kodiak Vicinity, Alaska. Center for Pacific Northwest Studies Occasional Paper 23. Western Washington University, Bellingham, 133 pp.", "Harmelin, J. - G., Ostrovsky, A. N., Caceres-Chamizo, J. P. & Sanner, J. (2011) Bryodiversity in the tropics: taxonomy of Microporella species (Bryozoa, Cheilostomata) with personate maternal zooids from Indian Ocean, Red Sea and southeast Mediterranean. Zootaxa, 2798 (1), 1 - 30. https: // doi. org / 10.11646 / zootaxa. 2798.1.1", "Suwa, T. & Mawatari, S. F. (1998) Revision of seven species of Microporella (Bryozoa, Cheilostomatida) from Hokkaido, Japan, using new taxonomic characters. Journal of Natural History, 32, 895 - 922. https: // doi. org / 10.1080 / 00222939800770461", "Liu, X., Yin, X. & Ma, J. (2001) Biology of Marine-Fouling Bryozoans in the Coastal Waters of China. Science Press, Beijing, 860 pp., 82 pls.", "Dick, M. H. & Grischenko, A. V. (2017) Rocky-intertidal cheilostome bryozoans from the vicinity of the Sesoko Biological Station, west-central Okinawa, Japan. Journal of Natural History, 51, 141 - 266. https: // doi. org / 10.1080 / 00222933.2016.1253797", "O'Dea, A. & Okamura, B. (1999) Influence of seasonal variation in temperature, salinity and food availability on module size and colony growth of the estuarine bryozoan Conopeum seurati. Marine Biology, 135, 581 - 588. https: // doi. org / 10.1007 / s 002270050659", "O'Dea, A. & Okamura, B. (2000) Intracolony variation in zooid size in cheilostome bryozoans as a new technique for investigation palaeoseasonality. Palaeogeography Palaeoclimatology Palaeoecology, 162, 319 - 332. https: // doi. org / 10.1016 / S 0031 - 0182 (00) 00136 - X"]}
format Text
author Dick, Matthew H.
Ngai, Nguyen Danh
Doan, Hung Dinh
author_facet Dick, Matthew H.
Ngai, Nguyen Danh
Doan, Hung Dinh
author_sort Dick, Matthew H.
title Microporella tonkinensis Dick & Ngai & Doan 2020, n. sp.
title_short Microporella tonkinensis Dick & Ngai & Doan 2020, n. sp.
title_full Microporella tonkinensis Dick & Ngai & Doan 2020, n. sp.
title_fullStr Microporella tonkinensis Dick & Ngai & Doan 2020, n. sp.
title_full_unstemmed Microporella tonkinensis Dick & Ngai & Doan 2020, n. sp.
title_sort microporella tonkinensis dick & ngai & doan 2020, n. sp.
publisher Zenodo
publishDate 2020
url https://dx.doi.org/10.5281/zenodo.3703680
https://zenodo.org/record/3703680
long_lat ENVELOPE(-55.200,-55.200,-63.200,-63.200)
ENVELOPE(-65.042,-65.042,-67.825,-67.825)
geographic Caceres
Indian
Pacific
Tonkin
geographic_facet Caceres
Indian
Pacific
Tonkin
genre Ketchikan
Kodiak
Alaska
genre_facet Ketchikan
Kodiak
Alaska
op_relation http://zenodo.org/record/3694740
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op_rights Open Access
info:eu-repo/semantics/openAccess
op_doi https://doi.org/10.5281/zenodo.3703680
https://doi.org/10.11646/zootaxa.4747.2.1
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spelling ftdatacite:10.5281/zenodo.3703680 2023-05-15T17:02:24+02:00 Microporella tonkinensis Dick & Ngai & Doan 2020, n. sp. Dick, Matthew H. Ngai, Nguyen Danh Doan, Hung Dinh 2020 https://dx.doi.org/10.5281/zenodo.3703680 https://zenodo.org/record/3703680 unknown Zenodo http://zenodo.org/record/3694740 http://publication.plazi.org/id/3104565CCC4FFFF6B106FFB1FFC61C7B http://zoobank.org/6AA8F5DC-8D70-42B0-B016-6F9C4211C471 https://zenodo.org/communities/biosyslit https://dx.doi.org/10.11646/zootaxa.4747.2.1 http://zenodo.org/record/3694740 http://publication.plazi.org/id/3104565CCC4FFFF6B106FFB1FFC61C7B https://dx.doi.org/10.5281/zenodo.3694764 http://zoobank.org/6AA8F5DC-8D70-42B0-B016-6F9C4211C471 https://dx.doi.org/10.5281/zenodo.3703679 https://zenodo.org/communities/biosyslit Open Access info:eu-repo/semantics/openAccess Biodiversity Taxonomy Animalia Bryozoa Gymnolaemata Cheilostomatida Microporellidae Microporella Microporella tonkinensis Taxonomic treatment article-journal Text ScholarlyArticle 2020 ftdatacite https://doi.org/10.5281/zenodo.3703680 https://doi.org/10.11646/zootaxa.4747.2.1 https://doi.org/10.5281/zenodo.3694764 https://doi.org/10.5281/zenodo.3703679 2022-02-08T12:04:14Z Microporella tonkinensis n. sp. (Fig. 12 A–D) urn:lsid:zoobank.org:act: D8480B51-CC0D-4B7B-B038-581C87933ADC Etymology. The specific name comes from the Gulf of Tonkin, wherein lies the Co To island group. Material examined. Holotype, VNMN-0248 (CT-6); paratype, VNMN-0249 (CT-41); both on SEM stubs. Measurements. Holotype , VNMN-0248: AzL , 0.43–0.59 (0.513 ± 0.050); AzW , 0.28–0.48 (0.357 ± 0.051); OrL , 0.062 –0.074 (0.068 ± 0.003); OrW , 0.106 –0.124 (0.113 ± 0.005); OvL , 0.19–0.23 (0.207 ± 0.011); OvW , 0.23–0.27 (0.260 ± 0.011); AvRL , 0.090 –0.114 (0.102 ± 0.007); AvRW , 0.046 –0.063 (0.053 ± 0.006) (n = 15, 1). Paratype , VNMN-0249: AzL , 0.34–0.49 (0.428 ± 0.039); AzW , 0.23–0.38 (0.294 ± 0.043); OrL , 0.056 –0.072 (0.066 ± 0.004); OrW , 0.087 –0.104 (0.098 ± 0.005); AvRL , 0.068 –0.092 (0.082 ± 0.007); AvRW , 0.038 –0.057 (0.046 ± 0.005) (n = 15, 1). Diagnosis. Orifice with straight, weakly denticulate proximal margin; small condyles mediad to proximolateral corners of orifice. Modal number of oral spines four or five; spine bases often evident in zooids away from colony margin. Ascopore non-cribriform, without conspicuous umbo; pustulation between ascopore and orifice. Avicularia lateral or proximolateral to ascopore, one per zooid, small relative to orifice size. Non-personate, non-ribbed ooecium retaining pair of spines at proximolateral corners. Description. Colony unilaminar, encrusting, sheet-like. Zooids irregularly hexagonal, distinct, delineated by groove. Frontal wall markedly convex, with two to five small, inconspicuous areolae along each lateral margin; perforated over entire surface by small pseudopores, except in zone between ascopore and orifice; texture uniformly coarsely pustulose between pores; pustules usually present between ascopore and orifice, sometimes in discrete row. Orifice broader than long, broadly D-shaped, surrounded by low, smooth rim; proximal margin straight or slightly concave between conspicuous triangular condyles; condyles positioned slightly mediad to proximolateral corners of orifice; proximal margin weakly denticulate. Oral spines short, articulated; three (n = 5) or four (n = 27) in colony CT-6; three (n = 1), four (n = 13), five (n = 17), or six (n = 4) in colony CT-41; spines retained even in older parts of colony. Ascopore small, broader than long, encircled by smooth, raised rim, separated from proximal orificial margin by 1–1.5 times height of ascopore plus rim; with small lingula projecting proximally, leaving C-shaped opening; margin of opening finely denticulate all around. Sometimes proximal part of ascopore rim widened and raised as low, smooth tubercle. One avicularium per zooid, lateral to ascopore and directed distolaterally or sometimes laterally, or along zooidal margin proximolateral to ascopore, sometimes at lateral corner of zooid or even more proximally, directed distolaterally or laterally. Frontal avicularia moderately small, rostrum raised, sides and edges smooth, crossbar complete, area proximal to crossbar semicircular, mandibular area triangular, slightly longer than broad, guttered distally to allow for setiform mandible; setiform portion of mandible as long as or shorter than triangular rostral portion. Ooecium prominent, resting on frontal wall of next-distal zooid, broader than long, closed by operculum, with narrow, smooth, flat zone around proximal margin where operculum meets ooecium; with many small pseudopores around steep periphery, few in central-proximal area; top more finely pustulose than frontal wall, coarsely pustulose around periphery, with some moderately large marginal openings at base around distal margin; ooecium appearing peripherally ribbed at low magnification. Most-proximal pair of spines lateral to orifice retained in ooecium-bearing zooids, abutting proximolateral corners of ooecium. Remarks. Microporella is a speciose genus, with roughly 140 mostly Recent species described worldwide, from polar regions to the tropics (Bock 2018). The generic features are rather stereotyped, simplifying identification to genus but complicating identification to species. Until relatively recently, taxonomists viewed the European species M. ciliata (Pallas, 1766) as having a cosmopolitan distribution that essentially mirrored the range of the genus. While many species worldwide were correctly delineated from M. ciliata on the basis of unique apomorphies or combinations of characters, in the older literature many specimens from the tropical Pacific and elsewhere more or less similar to M. ciliata were attributed to that species, without description; for example, Canu & Bassler (1929) list M. ciliata from Hong Kong and the Philippines. Some of the characters utilized in the taxonomy of this genus are variable rather than discrete. For example, M. germana Dick & Ross, 1988 at Kodiak, Alaska, had three to five oral spines, with the modal number four, but at Ketchikan had four to six spines, with the modal number five (Dick et al . 2005). Similar interpopulation variation is seen in the frequencies of zooids having single, paired, or no avicularia per zooid, and in the shape of the ascopore (e.g., Harmelin et al . 2011, p. 18, fig. 9). One useful character that distinguishes one large group of species from another is the presence or absence of a personate ooecium (ooecium-bearing zooids have a raised peristomial rim, proximal and lateral to the orifice, that meets the ooecium distally) (Harmelin et al . 2011); M. dentilingua above has a personate ooecium, whereas M. tonkinensis n. sp. does not. Several similar species in the northwestern Pacific differ from M. tonkinensis n. sp. in the following ways. Microporella formosa Suwa & Mawatari, 1998 from Hokkaido, Japan, has larger avicularia relative to orifice size; the proximal orificial margin is smooth, with the condyles larger and occupying the proximolateral corners of the orifice. The orifice in Microporella antiborealis Liu & Liu, 2001 [listed in Liu et al . (2001) as previously described rather than as a new species, but we could find no reference for Liu & Liu (2001)] from China has a similarly finely denticulate proximal margin, but has larger avicularia relative to orifice size, a larger ascopore, and modal spine number six, and lacks the pair of lateral oral spines remaining at the proximolateral corners of the ooecium. Like M. tonkinensis n. sp. , Microporella borealis Suwa & Mawatari, 1998 from Hokkaido, Japan, has a finely denticulate proximal orificial margin, and has four or five oral spines, retaining one pair at the proximolateral corners of the ooecium, but the condyles are larger and in the proximolateral corners of the orifice, the avicularia are much larger relative to orifice size, and there is no pustulation between the ascopore and orifice. Only two specimens of M. tonkinensis were found; these differed markedly in zooid size, with CT-6 having larger zooids than CT-41. This sort of variation seems to be common in tropical to subtropical cheilostomes, both within and between colonies; a similar case was observed in Rhynchozoon lunifrons Dick & Grischenko, 2017 at Okinawa. While some of this variation may be due to differences in temperature at time of growth (O’Dea & Okamura 1999, 2000), the degree of variation seems greater than occurs in cold-temperate faunas, and intercolony variation may require additional explanation. In any case, bryozoan taxonomists working in the tropics and subtropics should be aware that zooid size should be treated with caution as a taxonomic character. Distribution. Co To Island is the only known locality. : Published as part of Dick, Matthew H., Ngai, Nguyen Danh & Doan, Hung Dinh, 2020, Taxonomy and diversity of coelobite bryozoans from drift coral cobbles on Co To Island, northern Vietnam, pp. 201-252 in Zootaxa 4747 (2) on pages 227-228, DOI: 10.11646/zootaxa.4747.2.1, http://zenodo.org/record/3694740 : {"references": ["Bock, P. (2018) Indexes to bryozoan taxa. Available from: http: // www. bryozoa. net / indexes. html (accessed 15 December 2018)", "Pallas, P. S. (1766) Elenchus Zoophytorum Sistens Generum Adumbrationes Generaliores et Speciarum Cognitarum Cuccintas Descriptiones cum Selectis Auctorus Synonymis. Petrum van Cleef, Hagae-Comitum, 451 pp. https: // doi. org / 10.5962 / bhl. title. 6595", "Canu, F. & Bassler, R. S. (1929) Bryozoa of the Philippine region. United States National Museum Bulletin, 100, I-XI + 1 - 685.", "Dick, M. H. & Ross, J. R. P. (1988) Intertidal Bryozoa (Cheilostomata) of the Kodiak Vicinity, Alaska. Center for Pacific Northwest Studies Occasional Paper 23. Western Washington University, Bellingham, 133 pp.", "Harmelin, J. - G., Ostrovsky, A. N., Caceres-Chamizo, J. P. & Sanner, J. (2011) Bryodiversity in the tropics: taxonomy of Microporella species (Bryozoa, Cheilostomata) with personate maternal zooids from Indian Ocean, Red Sea and southeast Mediterranean. Zootaxa, 2798 (1), 1 - 30. https: // doi. org / 10.11646 / zootaxa. 2798.1.1", "Suwa, T. & Mawatari, S. F. (1998) Revision of seven species of Microporella (Bryozoa, Cheilostomatida) from Hokkaido, Japan, using new taxonomic characters. Journal of Natural History, 32, 895 - 922. https: // doi. org / 10.1080 / 00222939800770461", "Liu, X., Yin, X. & Ma, J. (2001) Biology of Marine-Fouling Bryozoans in the Coastal Waters of China. Science Press, Beijing, 860 pp., 82 pls.", "Dick, M. H. & Grischenko, A. V. (2017) Rocky-intertidal cheilostome bryozoans from the vicinity of the Sesoko Biological Station, west-central Okinawa, Japan. Journal of Natural History, 51, 141 - 266. https: // doi. org / 10.1080 / 00222933.2016.1253797", "O'Dea, A. & Okamura, B. (1999) Influence of seasonal variation in temperature, salinity and food availability on module size and colony growth of the estuarine bryozoan Conopeum seurati. Marine Biology, 135, 581 - 588. https: // doi. org / 10.1007 / s 002270050659", "O'Dea, A. & Okamura, B. (2000) Intracolony variation in zooid size in cheilostome bryozoans as a new technique for investigation palaeoseasonality. Palaeogeography Palaeoclimatology Palaeoecology, 162, 319 - 332. https: // doi. org / 10.1016 / S 0031 - 0182 (00) 00136 - X"]} Text Ketchikan Kodiak Alaska DataCite Metadata Store (German National Library of Science and Technology) Caceres ENVELOPE(-55.200,-55.200,-63.200,-63.200) Indian Pacific Tonkin ENVELOPE(-65.042,-65.042,-67.825,-67.825)