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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.860.34317" ID-GBIF-Dataset="e0a7142d-1eae-4254-b91a-30b70eb9b5ee" ID-PMC="PMC6690523" ID-Pensoft-Pub="1313-2970-860-183" ID-PubMed="31413657" ID-ZooBank="A3F9127D8ED24F8296A39510EB039A9C" ModsDocID="1313-2970-860-183" checkinTime="1565091121862" checkinUser="pensoft" docAuthor="Horvath, Elizabeth Anne" docDate="2019" docId="63F6A39DEE4EE27268338E6CC3127709" docLanguage="en" docName="ZooKeys 860: 183-306" docOrigin="ZooKeys 860" docSource="http://dx.doi.org/10.3897/zookeys.860.34317" docTitle="Plumarella longispina Kinoshita 1908" docType="treatment" docVersion="5" lastPageNumber="251" masterDocId="D577415D00092007FFDF4B2B606EFFFB" masterDocTitle="A review of gorgonian coral species (Cnidaria, Octocorallia, Alcyonacea) held in the Santa Barbara Museum of Natural History research collection: focus on species from Scleraxonia, Holaxonia, Calcaxonia - Part III: Suborder Holaxonia continued, and suborder Calcaxonia" masterLastPageNumber="306" masterPageNumber="183" pageNumber="248" updateTime="1668167482081" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>A review of gorgonian coral species (Cnidaria, Octocorallia, Alcyonacea) held in the Santa Barbara Museum of Natural History research collection: focus on species from Scleraxonia, Holaxonia, Calcaxonia - Part III: Suborder Holaxonia continued, and suborder Calcaxonia</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Horvath, Elizabeth Anne</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2019</mods:date>
<mods:detail type="volume">
<mods:number>860</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>183</mods:start>
<mods:end>306</mods:end>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.860.34317</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.860.34317</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-860-183</mods:identifier>
<mods:identifier type="ZooBank">A3F9127D8ED24F8296A39510EB039A9C</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="158525290" LSID="urn:lsid:plazi:treatment:63F6A39DEE4EE27268338E6CC3127709" httpUri="http://treatment.plazi.org/id/63F6A39DEE4EE27268338E6CC3127709" lastPageId="70" lastPageNumber="251" pageId="65" pageNumber="248">
<subSubSection pageId="65" pageNumber="248" type="nomenclature">
<paragraph pageId="65" pageNumber="248">
<taxonomicName LSID="63F6A39DEE4EE27268338E6CC3127709" authority="Kinoshita, 1908" authorityName="Kinoshita" authorityYear="1908" class="Anthozoa" family="Primnoidae" genus="Plumarella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plumarella longispina" order="Alcyonacea" pageId="65" pageNumber="248" phylum="Cnidaria" rank="species" species="longispina">Plumarella longispina Kinoshita, 1908</taxonomicName>
<figureCitation captionStart="Figure 39" captionStartId="F39" captionText="Figure 39. Plumarella longispina, SBMNH 422394. A Colony, 14 cm tall x 15 cm wide B Branch tips. Scale bar: 2 cm (A)." figureDoi="10.3897/zookeys.860.34317.figure39" httpUri="https://binary.pensoft.net/fig/314886" pageId="65" pageNumber="248">Figures 39A, B</figureCitation>
,
<figureCitation captionStart="Figure 40" captionStartId="F40" captionText="Figure 40. Plumarella longispina, SBMNH 422394, SEM image. A, B Body wall scales C Flatter coenenchymal scales D Marginal scales E Marginal spinous sclerites F-H Opercular scales (worn). Compare / contrast these images with those shown in Cairns 2011 (Figure 5, P. spicata)." figureDoi="10.3897/zookeys.860.34317.figure40" httpUri="https://binary.pensoft.net/fig/314887" pageId="65" pageNumber="248">
40
<normalizedToken originalValue="AH">A-H</normalizedToken>
</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="66" pageNumber="249" type="reference_group">
<paragraph pageId="66" pageNumber="249">
<taxonomicName class="Anthozoa" family="Primnoidae" genus="Plumarella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plumarella longispina" order="Alcyonacea" pageId="66" pageNumber="249" phylum="Cnidaria" rank="species" species="longispina">
<pageBreakToken pageId="66" pageNumber="249" start="start">Plumarella</pageBreakToken>
longispina
</taxonomicName>
Kinoshita, 1908a: 14, 15.
<bibRefCitation DOI="https://doi.org/10.5479/si.00963801.35-1658.681" author="Nutting, CC" journalOrPublisher="Proceedings of the United States National Museum" pageId="97" pageNumber="280" pagination="681 - 727" refId="B144" refString="Nutting, CC, 1909. Alcyonaria of the California coast. . Proceedings of the United States National Museum 35: 681 - 727" title="Alcyonaria of the California coast." url="https://doi.org/10.5479/si.00963801.35-1658.681" volume="35" year="1909">Nutting 1909</bibRefCitation>
: 716.
<bibRefCitation pageId="66" pageNumber="249" refId="B128">
<normalizedToken originalValue="Kükenthal">Kuekenthal</normalizedToken>
1924
</bibRefCitation>
: 260, 261.
</paragraph>
</subSubSection>
<subSubSection pageId="66" pageNumber="249" type="type locality">
<paragraph pageId="66" pageNumber="249">Type locality.</paragraph>
<paragraph pageId="66" pageNumber="249">N. Pacific Ocean, Japan, Honshu Island, Sagami Bay, Okinose Bank, 600 m.</paragraph>
</subSubSection>
<subSubSection pageId="66" pageNumber="249" type="type specimens">
<paragraph pageId="66" pageNumber="249">Type specimens.</paragraph>
<paragraph pageId="66" pageNumber="249">
<emphasis bold="true" pageId="66" pageNumber="249">Holotype</emphasis>
USNM 50117 [dry]; branch (from holotype), donated by Tokyo Imperial Museum; this material was examined. Main colony presumably still housed in collection at Tokyo Imperial Museum (all scientific and &quot;natural materials&quot; collections housed separately at what is now called the National Museum of Nature and Science); was unable to verify or confirm catalog number.
</paragraph>
</subSubSection>
<subSubSection pageId="66" pageNumber="249" type="materials_examined">
<paragraph pageId="66" pageNumber="249">Material examined.</paragraph>
<paragraph pageId="66" pageNumber="249">~33 lots (wet/dry) (see Appendix 3: List of material examined).</paragraph>
</subSubSection>
<subSubSection lastPageId="68" lastPageNumber="251" pageId="66" pageNumber="249" type="description">
<paragraph pageId="66" pageNumber="249">Description.</paragraph>
<paragraph lastPageId="68" lastPageNumber="251" pageId="66" pageNumber="249">
<emphasis italics="true" pageId="66" pageNumber="249">Colony</emphasis>
(
<figureCitation captionStart="Figure 39" captionStartId="F39" captionText="Figure 39. Plumarella longispina, SBMNH 422394. A Colony, 14 cm tall x 15 cm wide B Branch tips. Scale bar: 2 cm (A)." figureDoi="10.3897/zookeys.860.34317.figure39" httpUri="https://binary.pensoft.net/fig/314886" pageId="66" pageNumber="249">Figure 39A</figureCitation>
) exhibits dense, alternate, pinnate branching in one plane, leading to flabellate form. Main stem somewhat flattened, giving rise to alternate main branches at irregular distances; both main stem and branches may subdivide. Each main branch gives forth regularly alternate, slightly smaller branches that do not subdivide. Branchlets flattened, 1.5 mm thick (
<figureCitation captionStart="Figure 39" captionStartId="F39" captionText="Figure 39. Plumarella longispina, SBMNH 422394. A Colony, 14 cm tall x 15 cm wide B Branch tips. Scale bar: 2 cm (A)." figureDoi="10.3897/zookeys.860.34317.figure39" httpUri="https://binary.pensoft.net/fig/314886" pageId="66" pageNumber="249">Figure 39B</figureCitation>
). Polyps small, short, cylindrical projections, 0.5 mm tall (to summit of operculum), 0.5 mm across, 1.5 mm apart; arranged laterally in two opposite rows on flattened stems, branches and branchlets; some polyps placed such that they project toward a front side of colony, with back of colony smooth; strictly alternate to strictly opposite in different parts of colony, with upper edge of one polyp ordinarily reaching to base of next one above. Polyp aperture pointed upward, slightly outward. Walls of polyps armed with sclerites; these conspicuous, flattened scales, vary greatly in size and form in different polyps. Color of colony (? alive) generally white; dry or in alcohol, dull creamy-white; some preserved colonies light grayish-brown, with surfaces of stem and branches being more distinctly gray. Sclerites (
<figureCitation captionStart="Figure 40" captionStartId="F40" captionText="Figure 40. Plumarella longispina, SBMNH 422394, SEM image. A, B Body wall scales C Flatter coenenchymal scales D Marginal scales E Marginal spinous sclerites F-H Opercular scales (worn). Compare / contrast these images with those shown in Cairns 2011 (Figure 5, P. spicata)." figureDoi="10.3897/zookeys.860.34317.figure40" httpUri="https://binary.pensoft.net/fig/314887" pageId="66" pageNumber="249">
Figure 40
<normalizedToken originalValue="AH">A-H</normalizedToken>
</figureCitation>
) quite varied in form, generally more or less flattened into scales; thin, cycloid. Key characteristic sclerite a flattened basal portion bearing on its distal edge long thorn-like processes (spines) projecting above margin of polyp (
<figureCitation captionStart="Figure 40" captionStartId="F40" captionText="Figure 40. Plumarella longispina, SBMNH 422394, SEM image. A, B Body wall scales C Flatter coenenchymal scales D Marginal scales E Marginal spinous sclerites F-H Opercular scales (worn). Compare / contrast these images with those shown in Cairns 2011 (Figure 5, P. spicata)." figureDoi="10.3897/zookeys.860.34317.figure40" httpUri="https://binary.pensoft.net/fig/314887" pageId="66" pageNumber="249">Figure 40E</figureCitation>
). Many scales ornamented with convex, ctenate margin. Surfaces of scales ornamented with evenly, closely distributed granules, irregularly placed nodular warts and occasional spines. Typical arrangement of scales on polyp wall is eight lon
<pageBreakToken pageId="67" pageNumber="250" start="start">gitudinal</pageBreakToken>
rows, each row having roughly four scales in a ring; two proximal rings composed of broad curved scales with their distal convex edges ctenate, distal-most marginal ring composed of scales (with no keel), bearing prominent thorn-like, unwarted spines extending beyond end of operculum. Marginal spines usually number from two to six, two of which (abaxial) are often distinctly longer than the others. Operculum composed of eight irregularly shaped scales, not keeled, points of which often joined into spine-like processes (
<figureCitation captionStart="Figure 40" captionStartId="F40" captionText="Figure 40. Plumarella longispina, SBMNH 422394, SEM image. A, B Body wall scales C Flatter coenenchymal scales D Marginal scales E Marginal spinous sclerites F-H Opercular scales (worn). Compare / contrast these images with those shown in Cairns 2011 (Figure 5, P. spicata)." figureDoi="10.3897/zookeys.860.34317.figure40" httpUri="https://binary.pensoft.net/fig/314887" pageId="67" pageNumber="250">
Figure 40
<normalizedToken originalValue="FH">F-H</normalizedToken>
</figureCitation>
). Adcauline opercular scales reduced to nar
<pageBreakToken pageId="68" pageNumber="251" start="start">row</pageBreakToken>
band, the antero-lateral processes from proximal rings of sclerites being the only ones that meet to complete the ring on abcauline side.
</paragraph>
<caption doi="10.3897/zookeys.860.34317.figure39" httpUri="https://binary.pensoft.net/fig/314886" pageId="68" pageNumber="251" start="Figure 39" startId="F39">
<paragraph pageId="68" pageNumber="251">
Figure 39.
<taxonomicName class="Anthozoa" family="Primnoidae" genus="Plumarella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plumarella longispina" order="Alcyonacea" pageId="68" pageNumber="251" phylum="Cnidaria" rank="species" species="longispina">
<emphasis italics="true" pageId="68" pageNumber="251">Plumarella longispina</emphasis>
</taxonomicName>
, SBMNH 422394.
<emphasis bold="true" pageId="68" pageNumber="251">A</emphasis>
Colony, 14 cm tall
<normalizedToken originalValue="×">x</normalizedToken>
15 cm wide
<emphasis bold="true" pageId="68" pageNumber="251">B</emphasis>
Branch tips. Scale bar: 2 cm (
<emphasis bold="true" pageId="68" pageNumber="251">A</emphasis>
).
</paragraph>
</caption>
<caption doi="10.3897/zookeys.860.34317.figure40" httpUri="https://binary.pensoft.net/fig/314887" pageId="68" pageNumber="251" start="Figure 40" startId="F40">
<paragraph pageId="68" pageNumber="251">
Figure 40.
<taxonomicName class="Anthozoa" family="Primnoidae" genus="Plumarella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plumarella longispina" order="Alcyonacea" pageId="68" pageNumber="251" phylum="Cnidaria" rank="species" species="longispina">
<emphasis italics="true" pageId="68" pageNumber="251">Plumarella longispina</emphasis>
</taxonomicName>
, SBMNH 422394, SEM image.
<emphasis bold="true" pageId="68" pageNumber="251">A, B</emphasis>
Body wall scales
<emphasis bold="true" pageId="68" pageNumber="251">C</emphasis>
Flatter coenenchymal scales
<emphasis bold="true" pageId="68" pageNumber="251">D</emphasis>
Marginal scales
<emphasis bold="true" pageId="68" pageNumber="251">E</emphasis>
Marginal spinous sclerites
<emphasis bold="true" pageId="68" pageNumber="251">
<normalizedToken originalValue="FH">F-H</normalizedToken>
</emphasis>
Opercular scales (worn). Compare/contrast these images with those shown in
<bibRefCitation DOI="https://doi.org/10.5479/si.00810282.634" author="Cairns, SD" journalOrPublisher="Pacific Science" pageId="89" pageNumber="272" refId="B34" refString="Cairns, SD, 2011. . https://doi.org/10.5479/si.00810282.634" url="https://doi.org/10.5479/si.00810282.634" year="2011">Cairns 2011</bibRefCitation>
(
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Swiftia pacifica, specimen 41 - 39 - 1 (Alaska Fisheries Service, Gulf of Alaska); looking in same form of that seen in Figure 3, light microscopy arrays. A (4 x) showing variety of sclerites, particularly the characteristic &quot; fingerbiscuit-rod &quot; seen in the genus Swiftia. Sclerites from specimen examined for Bob Stone, Alaska Fisheries Service B Higher magnification, 10 x, showing all sclerite forms, including obvious anthocodial fingerbiscuit-rods. The larger spindles measure ~ 300 µm long, smaller spindles of ~ 200 µm, and the rods range from 308 - 370 µm in length." figureDoi="10.3897/zookeys.860.34317.figure5" httpUri="https://binary.pensoft.net/fig/314852" pageId="68" pageNumber="251">Figure 5</figureCitation>
,
<taxonomicName lsidName="P. spicata" pageId="68" pageNumber="251" rank="species" species="spicata">
<emphasis italics="true" pageId="68" pageNumber="251">P. spicata</emphasis>
</taxonomicName>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="68" pageNumber="251" type="etymology">
<paragraph pageId="68" pageNumber="251">Etymology.</paragraph>
<paragraph pageId="68" pageNumber="251">
From the Latin,
<emphasis italics="true" pageId="68" pageNumber="251">longi</emphasis>
- = long and
<emphasis italics="true" pageId="68" pageNumber="251">spina</emphasis>
- = spine; long-spined, referencing the spinose marginal sclerites that extend beyond end of operculum on polyps.
</paragraph>
</subSubSection>
<subSubSection pageId="68" pageNumber="251" type="distribution">
<paragraph pageId="68" pageNumber="251">Distribution.</paragraph>
<paragraph pageId="68" pageNumber="251">
Found off California coast between ~55-735 m. Of specimens examined, could not confirm that this species is found off the Oregon coast (thus far, all specimens examined were collected either from Baja California [Mexico] and California [USA] or Washington [USA]; it seems odd that it would skip an entire area between CA and WA). Based on material collected by staff of Olympic Coast National Marine Sanctuary (May, 2006 and July, 2008) that was examined, seen off northwest Washington coast at depths of at least ~208-309 m. Specimens from the genus have been taken in Alaskan waters (Bering Sea, etc.) in depths from 85-2514 m; collection data for these specimens can be found by doing a search of the online data base for the NMNH, Smithsonian, Invertebrate Collection. Listings of this particular species (by
<bibRefCitation author="Wing, BL" journalOrPublisher="NOAA Technical Memoires, NMFS-AFSC- 146" pageId="98" pageNumber="281" publicationUrl="www.afsc.noaa.gov/publications/afsc-tm/noaa-tm-afsc-146.pdf" refId="B188" refString="Wing, BL, Barnard, DR, 2004. A field Guide to Alaskan Corals. . NOAA Technical Memoires, NMFS-AFSC-146" title="A field Guide to Alaskan Corals." url="www.afsc.noaa.gov/publications/afsc-tm/noaa-tm-afsc-146.pdf" year="2004">Wing and Barnard 2004</bibRefCitation>
;
<bibRefCitation DOI="https://doi.org/10.1111/j.1365-2419.2005.00371.x" author="Heifetz, J" journalOrPublisher="Hydrobiologia" pageId="94" pageNumber="277" refId="B106" refString="Heifetz, J, Wing, BL, Stone, RP, Malacha, PW, Courtney, DL, 2005. . https://doi.org/10.1111/j.1365-2419.2005.00371.x" url="https://doi.org/10.1111/j.1365-2419.2005.00371.x" year="2005">Heifetz et. al. 2005</bibRefCitation>
; and
<bibRefCitation pageId="68" pageNumber="251" refId="B157">Stone and Shotwell 2007</bibRefCitation>
) mentioned in Cairns, 2011, could not be confirmed.
</paragraph>
</subSubSection>
<subSubSection pageId="68" pageNumber="251" type="biology_ecology">
<paragraph pageId="68" pageNumber="251">Biology.</paragraph>
<paragraph pageId="68" pageNumber="251">
Work by
<bibRefCitation author="Lissner, AL" journalOrPublisher="Bulletin of the Southern California Academy of Sciences" pageId="96" pageNumber="279" pagination="87 - 101" refId="B134" refString="Lissner, AL, Dorsey, JH, 1986. Deep-water biological assemblages of a hard-bottom bank-ridge complex of the Southern California Continental Borderland. . Bulletin of the Southern California Academy of Sciences 85: 87 - 101" title="Deep-water biological assemblages of a hard-bottom bank-ridge complex of the Southern California Continental Borderland." volume="85" year="1986">Lissner and Dorsey (1986)</bibRefCitation>
along Tanner and Cortes Banks and the Santa Rosa-Cortes Ridge area off southern California showed a depth range as follows: at depths &lt;67 m the species is sparse, at depths ranging from 67-122 m the species is common to abundant, and at depths below 122 m, again becoming sparse. Deep-water video images taken by MBARI indicated the possibility of the genus (perhaps this species) being more common at greater depth (at least in some areas) than once thought.
</paragraph>
<paragraph pageId="68" pageNumber="251">In all specimens examined, only one had any other organism associated with it; on this specimen there appeared two anemones, both on branches near the tip. One, the larger of the two, is on the exposed axis. On this same specimen, on the area of branches just above the base, there appeared to be the anchor tendrils from the egg case of a shark. These tendrils are quite thin, but with the stiff curl they usually display. Egg cases were noticeable on specimens collected by OCNMS in May 2006. Colonies of this species are quite rigid, so it is likely that they provide good anchorage.</paragraph>
</subSubSection>
<subSubSection lastPageId="70" lastPageNumber="253" pageId="68" pageNumber="251" type="remarks">
<paragraph pageId="68" pageNumber="251">Remarks.</paragraph>
<paragraph pageId="68" pageNumber="251">
A key data point in the distribution of this species was
<normalizedToken originalValue="Nuttings">Nutting's</normalizedToken>
specimen locality (1909):
<normalizedToken originalValue="Albatross">'Albatross'</normalizedToken>
station 4359, Point Loma light-house,
<geoCoordinate degrees="32" direction="north" minutes="42" orientation="latitude" precision="15" seconds="00" value="32.7">32°42'00&quot;N</geoCoordinate>
,
<geoCoordinate degrees="117" direction="west" minutes="14" orientation="longitude" precision="15" seconds="00" value="-117.23333">117°14'00&quot;W</geoCoordinate>
(N 85, E 9 miles), 191 fathoms (347 m). This specimen currently housed at NMNH (USNM 25429); specimen was examined.
</paragraph>
<paragraph pageId="68" pageNumber="251">
In a comparison with a different species (from the Aleutian Islands,
<taxonomicName class="Anthozoa" family="Primnoidae" genus="Plumarella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Plumarella spicata" order="Alcyonacea" pageId="68" pageNumber="251" phylum="Cnidaria" rank="species" species="spicata">
<emphasis italics="true" pageId="68" pageNumber="251">Plumarella spicata</emphasis>
</taxonomicName>
Nutting, 1912), it presented marginal scales that were similar in shape to those seen in this species, but the spinous process of the marginal scales in
<taxonomicName lsidName="P. longispina" pageId="68" pageNumber="251" rank="species" species="longispina">
<emphasis italics="true" pageId="68" pageNumber="251">P. longispina</emphasis>
</taxonomicName>
are much less ornamented. As well, all of the operculars in the species described here display areas of surface that appear very smooth and undecorated; in
<taxonomicName lsidName="P. spicata" pageId="68" pageNumber="251" rank="species" species="spicata">
<emphasis italics="true" pageId="68" pageNumber="251">P. spicata</emphasis>
</taxonomicName>
, surface ornamentation is more prevalent, although perhaps not continuous along entire surface. Colony form of
<taxonomicName lsidName="P. spicata" pageId="68" pageNumber="251" rank="species" species="spicata">
<emphasis italics="true" pageId="68" pageNumber="251">P. spicata</emphasis>
</taxonomicName>
(delicate and flimsy, more or less dichotomously branched), does not match what is seen for this species.
</paragraph>
<paragraph lastPageId="70" lastPageNumber="253" pageId="68" pageNumber="251">
Unless there are very subtle differences, e.g., characteristics that might specify several subspecies, this species seemed to be one of the most abundant deep-water prim
<pageBreakToken pageId="69" pageNumber="252" start="start">noids</pageBreakToken>
occurring in the California Bight (and elsewhere). Its overall colony form is quite distinctive, and easily recognizable. While appearing to be quite delicate, closer examination and handling indicated that it is actually fairly hardy. In the near future, an examination of all specimens in the SBMNH collection will have to be undertaken, with special attention paid to any feature(s) that could be assessed as a key characteris
<pageBreakToken pageId="70" pageNumber="253" start="start">tic</pageBreakToken>
that might show some degree of variability. The question arose as to whether there are transitional variations over the entire range of this species, and if so, whether those variations might subdivide the specimens, such that they point in the direction of distinct subspecies (or for that matter, species). Molecular studies on any of those groupings could add further clarity. However, it may be that this is simply an enormously successful species, thus very common, with adequate and successful dispersal abilities. Cordeiro et al. (2019) shows
<taxonomicName lsidName="P. longispina" pageId="70" pageNumber="253" rank="species" species="longispina">
<emphasis italics="true" pageId="70" pageNumber="253">P. longispina</emphasis>
</taxonomicName>
with accepted species status.
</paragraph>
</subSubSection>
</treatment>
</document>