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<mods:title id="AD0716EEA926326A0DA53C7D3F81CB60">Sperm ultrastructure in Hemidonax pictus (Hemidonacidae, Bivalvia, Mollusca): comparison with other heterodonts, especially Cardiidae, Donacidae and Crassatelloidea</mods:title>
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<paragraph id="9B0E36F4633F1E2D18594A1CFC3FFD85" blockId="10.[823,1404,585,606]" box="[823,1404,585,606]" pageId="10" pageNumber="335">
<heading id="C0468198633F1E2D18594A1CFC3FFD85" box="[823,1404,585,606]" centered="true" fontSize="9" level="2" pageId="10" pageNumber="335" reason="2">
COMPARISON OF
<taxonomicName id="5CB14D77633F1E2D18924A1CFDC5FD85" ID-CoL="7P4ZL" authorityName="Morch" authorityYear="1778" box="[1020,1158,585,606]" class="Bivalvia" family="Hemidonacidae" genus="Hemidonax" kingdom="Animalia" order="Venerida" pageId="10" pageNumber="335" phylum="Mollusca" rank="genus">
<emphasis id="A9C5EAE6633F1E2D18924A1CFDC5FD85" box="[1020,1158,585,606]" italics="true" pageId="10" pageNumber="335">HEMIDONAX</emphasis>
</taxonomicName>
SPERM MORPHOLOGY
</heading>
</paragraph>
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<subSubSection id="D3AB657F633F1E2C18E74A38FAD3F8B7" lastPageId="11" lastPageNumber="336" pageId="10" pageNumber="335" type="description">
<paragraph id="9B0E36F4633F1E2D18E74A38FC69FDA5" blockId="10.[905,1322,620,637]" box="[905,1322,620,637]" pageId="10" pageNumber="335">WITH OTHER HETERODONT BIVALVES</paragraph>
<paragraph id="9B0E36F4633F1E2D18484ADAFD6BFA18" blockId="10.[806,1422,655,1901]" pageId="10" pageNumber="335">
Spermatozoa of
<taxonomicName id="5CB14D77633F1E2D188D4ADAFDF7FD7D" baseAuthorityName="Tryon" baseAuthorityYear="1870" box="[995,1204,655,677]" class="Bivalvia" family="Hemidonacidae" genus="Hemidonax" kingdom="Animalia" order="Venerida" pageId="10" pageNumber="335" phylum="Mollusca" rank="species" species="pictus">
<emphasis id="A9C5EAE6633F1E2D188D4ADAFDF7FD7D" box="[995,1204,655,677]" italics="true" pageId="10" pageNumber="335">Hemidonax pictus</emphasis>
</taxonomicName>
are of the simple, aquasperm
<typeStatus id="440A8856633F1E2D18DD4AFBFAA7FD1B" box="[947,996,686,707]" pageId="10" pageNumber="335">type</typeStatus>
characteristic of molluscs showing aquatic fertilization, especially the
<taxonomicName id="5CB14D77633F1E2D1FD74A99FC5BFD3A" authorityName="Linnaeus" authorityYear="1758" box="[1209,1304,716,738]" class="Bivalvia" kingdom="Animalia" pageId="10" pageNumber="335" phylum="Mollusca" rank="class">Bivalvia</taxonomicName>
(
<bibRefCitation id="FF204B05633F1E2D1E494A99FA1CFCD9" author="Franzen A" pageId="10" pageNumber="335" pagination="399 - 456" refId="ref11484" refString="Franzen A. 1955. Comparative morphological investigations into the spermiogenesis among Mollusca. Zoologiska Bidrag fran Uppsala 30: 399 - 456." type="journal article" year="1955">Franzén, 1955</bibRefCitation>
,
<bibRefCitation id="FF204B05633F1E2D18184ABEFAECFCD8" author="Franzen A" box="[886,943,747,768]" pageId="10" pageNumber="335" pagination="199 - 214" refId="ref11539" refString="Franzen A. 1983. Ultrastructural studies of spermatozoa in three bivalve species with notes on evolution of elongated sperm nucleus in primitive spermatozoa. Gamete Research 7: 199 - 214." type="journal article" year="1983">1983</bibRefCitation>
;
<bibRefCitation id="FF204B05633F1E2D18AB4ABEFD37FCD8" author="Popham JD" box="[965,1140,747,769]" pageId="10" pageNumber="335" pagination="1 - 20" refId="ref12929" refString="Popham JD. 1979. Comparative spermatozoon morphology and bivalve phylogeny. Malacological Review 12: 1 - 20." type="journal article" year="1979">Popham, 1979</bibRefCitation>
;
<bibRefCitation id="FF204B05633F1E2D1FE44ABEFCCBFCD8" author="Hodgson AN &amp; Bernard RTF &amp; Van der Horst G" box="[1162,1416,747,769]" pageId="10" pageNumber="335" pagination="257 - 265" refId="ref12075" refString="Hodgson AN, Bernard RTF, Van der Horst G. 1990. Comparative spermatology of three species of Donax (Bivalvia) from South Africa. Journal of Molluscan Studies 56: 257 - 265." type="journal article" year="1990">
Hodgson
<emphasis id="A9C5EAE6633F1E2D1F914AB9FC73FCD8" box="[1279,1328,747,769]" italics="true" pageId="10" pageNumber="335">et al</emphasis>
., 1990
</bibRefCitation>
;
<bibRefCitation id="FF204B05633F1E2D18484B5FFA81FCC7" author="Healy JM" box="[806,962,777,800]" pageId="10" pageNumber="335" pagination="99 - 113" refId="ref11760" refString="Healy JM. 1996 a. Molluscan sperm ultrastructure: correlation with taxonomic units within the Gastropoda, Cephalopoda and Bivalvia. In: Taylor J, ed. Origin and evolutionary radiation of the Mollusca. Oxford: Oxford University Press, 99 - 113." type="book chapter" year="1996">Healy, 1996a</bibRefCitation>
), and also seen in a number of other invertebrate groups (e.g. cnidarians, polychaete annelids, brachiopods, echiurids, sipunculans
<bibRefCitation id="FF204B05633F1E2D1E494B12FA1CFCA3" author="Franzen A" pageId="10" pageNumber="335" pagination="355 - 482" refId="ref11508" refString="Franzen A. 1956. On spermiogenesis, morphology of the spermatozoon, and the biology of fertilization among invertebrates. Zoologiska Bidrag fran Uppsala 31: 355 - 482." type="journal article" year="1956">Franzén, 1956</bibRefCitation>
;
<bibRefCitation id="FF204B05633F1E2D18014B30FDD8FCA3" author="Baccetti B &amp; Afzelius BA" box="[879,1179,869,891]" pageId="10" pageNumber="335" refId="ref11254" refString="Baccetti B, Afzelius BA. 1976. Biology of the sperm cell. Monographs in Developmental Biology. Basel: S. Karger." type="book" year="1976">Baccetti &amp; Afzelius, 1976</bibRefCitation>
;
<bibRefCitation id="FF204B05633F1E2D1FC54B30FA23FC42" author="Jamieson BGM &amp; Rouse GW" pageId="10" pageNumber="335" pagination="93 - 157" refId="ref12212" refString="Jamieson BGM, Rouse GW. 1989. The spermatozoa of the Polychaeta (Annelida): an ultrastructural view. Biological Reviews 64: 93 - 157." type="journal article" year="1989">Jamieson &amp; Rouse, 1989</bibRefCitation>
).
<bibRefCitation id="FF204B05633F1E2D18EA4BD1FD99FC42" author="Rouse GW &amp; Jamieson BGM" box="[900,1242,900,922]" pageId="10" pageNumber="335" pagination="573 - 584" refId="ref13015" refString="Rouse GW, Jamieson BGM. 1987. An ultrastructural study of the spermatozoa of the polychaetes Eurythoe complanata (Amphinomidae), Clymenella laseroni and Micromaldane laseroni (Maldanidae) with definition of sperm types in relation to reproductive biology. Journal of Submicroscopic Cytology 19: 573 - 584." type="journal article" year="1987">Rouse &amp; Jamieson (1987)</bibRefCitation>
differentiated aquasperm into those that fertilize eggs in the ambient water (ect-aquasperm) and those that fertilize aquatically but within a protected space such as a worm tube or a molluscan mantle cavity (ent-aquasperm). In the absence of any information on the fertilization biology of
<taxonomicName id="5CB14D77633F1E2D1FE34C69FDB0FB89" baseAuthorityName="Tryon" baseAuthorityYear="1870" box="[1165,1267,1084,1105]" class="Bivalvia" family="Hemidonacidae" genus="Hemidonax" kingdom="Animalia" order="Venerida" pageId="10" pageNumber="335" phylum="Mollusca" rank="species" species="pictus">
<emphasis id="A9C5EAE6633F1E2D1FE34C69FDB0FB89" box="[1165,1267,1084,1105]" italics="true" pageId="10" pageNumber="335">H. pictus</emphasis>
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, it is impossible to characterize the
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of aquasperm in this species. However it is worth noting that in shipworms (
<taxonomicName id="5CB14D77633F1E2D18404CC2FD13FB75" authority=", Pholadoidea" authorityName="Pholadoidea" box="[814,1104,1175,1197]" class="Bivalvia" family="Teredinidae" kingdom="Animalia" order="Myida" pageId="10" pageNumber="335" phylum="Mollusca" rank="family">Teredinidae, Pholadoidea</taxonomicName>
)
<bibRefCitation id="FF204B05633F1E2D1F0E4CC2FC4EFB75" author="Popham JD" box="[1120,1293,1175,1197]" pageId="10" pageNumber="335" pagination="291 - 297" refId="ref12887" refString="Popham JD. 1974. Comparative morphometrics of the acrosomes of the sperms of ' externally' and ' internally' fertilizing sperms of the shipworms (Teredinidae, Bivalvia, Mollusca). Cell and Tissue Research 150: 291 - 297." type="journal article" year="1974">Popham (1974)</bibRefCitation>
found that species fertilizing within the mantle cavity had smaller acrosomes than those that fertilized externally (i.e. in the ambient seawater). If such a correlation holds among the
<taxonomicName id="5CB14D77633F1E2D1F524D47FDD8FAF0" authorityName="Linnaeus" authorityYear="1758" box="[1084,1179,1298,1320]" class="Bivalvia" kingdom="Animalia" pageId="10" pageNumber="335" phylum="Mollusca" rank="class">Bivalvia</taxonomicName>
in general, then the small size of the acrosome of
<taxonomicName id="5CB14D77633F1E2D1F184D64FD9FFA9E" baseAuthorityName="Tryon" baseAuthorityYear="1870" box="[1142,1244,1329,1350]" class="Bivalvia" family="Hemidonacidae" genus="Hemidonax" kingdom="Animalia" order="Venerida" pageId="10" pageNumber="335" phylum="Mollusca" rank="species" species="pictus">
<emphasis id="A9C5EAE6633F1E2D1F184D64FD9FFA9E" box="[1142,1244,1329,1350]" italics="true" pageId="10" pageNumber="335">H. pictus</emphasis>
</taxonomicName>
compared with most investigated heterodonts (see
<figureCitation id="038A2A71633F1E2D1FA04D1AFC57FABC" box="[1230,1300,1359,1381]" captionStart="Figure 8" captionStartId="18.[144,223,198,217]" captionText="Figure 8. Diagram showing comparative sperm ultrastructure (TEM) (longitudinal sections through head, midpiece and proximal portion of flagellum) of Hemidonax pictus (Tryon, 1870) (A), Cardiidae (BK), Donacidae (LP) and selected Veneridae (QV). BK, Cardiidae. B, Fragum unedo (Linnaeus, 1758); C, Lunulicardia hemicardium (Linnaeus, 1758); D, Vasticardium vertebratum (Jonas, 1844); E, Papyridea semisulcata (Gray, 1825); F, Fulvia tenuicostata (Lamarck, 1819); G, H, Cerastoderma spp.; I, Hippopus hippopus (Linnaeus, 1758); J, Tridacna gigas (Linnaeus, 1758); K, Tridacna maxima (Röding, 1798). LP, Donacidae. L, Donax deltoides Lamarck, 1818; M, Donax madagascariensis Wood, 1828; N, Donax sordidus Hanley, 1845; O, Donax serra Röding, 1798; P, Donax trunculus Linnaeus, 1758. QV, Veneridae. Q, Venerupis aurea (Gmelin, 1791); R, Venerupis corrugata (Gmelin, 1791); S, Petricola lapicida (Gmelin, 1791); T, Antigona chemnitzii (Hanley, 1844); U, Dosinia nedigna (Iredale, 1930); V, Lioconcha annettae Lamprell &amp; Whitehead, 1990. Scale bar for Hemidonax pictus (= 1.0 Mm) applies to all other figures except H (= 2.0 Mm). Sources of data: AE, L, SV (this paper; J. M. Healy, P. M. Mikkelsen &amp; R. Bieler, unpubl. data); F (Popham, 1979); G, H [composite reconstruction based on Cerastoderma lamarcki (Reeve, 1844) from Drozdov et al., 2001 and Cerastoderma edule (Linnaeus, 1758) from Sousa &amp; Azevedo, 1988 and Sousa et al., 1998]; IK (Keys &amp; Healy, 1999, 2000); MO (Hodgson et al., 1990); P (Sousa &amp; Oliveira, 1994); Q, R (Pochon-Masson &amp; Gharagozlou, 1970; Gharagozlou-Van Ginneken &amp; Pochon-Masson, 1971)." pageId="10" pageNumber="335">Figs 8</figureCitation>
,
<figureCitation id="038A2A71633F1E2D1E484D1AFC77FABD" box="[1318,1332,1359,1381]" captionStart="Figure 9" captionStartId="20.[144,223,198,217]" captionText="Figure 9. Diagram showing comparative acrosomal complex ultrastructure (TEM) of Hemidonax pictus (A), Cardiidae, Donacidae and selected Veneridae. BH, Cardiidae: B, Fragum unedo; C, Vasticardium vertebratum; D, Lunulicardia hemicardium; E, Fulvia tenuicostata; F, Papyridea semisulcata; G, Tridacna maxima; H, Hippopus hippopus. IL, Donacidae: I, Donax deltoides; J, Donax madagascariensis; K, Donax serra; L, Donax trunculus. MP, Veneridae: M, Lioconcha annettae; N, Dosinia nedigna; O, Antigona chemnitzii; P, Venerupis corrugata. Scale bar (1.0 Mm) applies to all figures. Sources of data: AD, F, I, MO (this paper; J. M. Healy, P. M. Mikkelsen &amp; R. Bieler, unpubl. data); E (Popham, 1979); G, H (Keys &amp; Healy, 1999, 2000); P (Gharagozlou-Van Ginneken &amp; Pochon-Masson, 1971); J, K (Hodgson et al., 1990); L (Sousa &amp; Oliveira, 1994)." pageId="10" pageNumber="335">9</figureCitation>
) would suggest the likelihood of ent-aquatic (mantle cavity) fertilization in this species and probably in other species of
<taxonomicName id="5CB14D77633F1E2D18F14DFEFD62FA18" authorityName="Morch" authorityYear="1778" box="[927,1057,1451,1472]" class="Bivalvia" family="Hemidonacidae" genus="Hemidonax" kingdom="Animalia" order="Venerida" pageId="10" pageNumber="335" phylum="Mollusca" rank="genus">
<emphasis id="A9C5EAE6633F1E2D18F14DFEFD62FA18" box="[927,1057,1451,1472]" italics="true" pageId="10" pageNumber="335">Hemidonax</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="9B0E36F4633F1E2C18514D9CFAD3F8B7" blockId="10.[806,1422,655,1901]" lastBlockId="11.[826,1441,1789,1903]" lastPageId="11" lastPageNumber="336" pageId="10" pageNumber="335">
Whereas it is true to state that spermatozoa of
<taxonomicName id="5CB14D77633F1E2D18484DBDFAB4FA26" baseAuthorityName="Tryon" baseAuthorityYear="1870" box="[806,1015,1512,1534]" class="Bivalvia" family="Hemidonacidae" genus="Hemidonax" kingdom="Animalia" order="Venerida" pageId="10" pageNumber="335" phylum="Mollusca" rank="species" species="pictus">
<emphasis id="A9C5EAE6633F1E2D18484DBDFAB4FA26" box="[806,1015,1512,1534]" italics="true" pageId="10" pageNumber="335">Hemidonax pictus</emphasis>
</taxonomicName>
do not exhibit any unique or new features, the precise combination of features is distinctive and presumably characteristic of the genus as a whole. Like other heterodonts,
<taxonomicName id="5CB14D77633F1E2D1FB84E11FC7EF981" baseAuthorityName="Tryon" baseAuthorityYear="1870" box="[1238,1341,1604,1625]" class="Bivalvia" family="Hemidonacidae" genus="Hemidonax" kingdom="Animalia" order="Venerida" pageId="10" pageNumber="335" phylum="Mollusca" rank="species" species="pictus">
<emphasis id="A9C5EAE6633F1E2D1FB84E11FC7EF981" box="[1238,1341,1604,1625]" italics="true" pageId="10" pageNumber="335">H. pictus</emphasis>
</taxonomicName>
shows a well-developed basal ring component of the acrosomal vesicle (see
<figureCitation id="038A2A71633F1E2D18804ED4FD73F94E" box="[1006,1072,1665,1687]" captionStart="Figure 8" captionStartId="18.[144,223,198,217]" captionText="Figure 8. Diagram showing comparative sperm ultrastructure (TEM) (longitudinal sections through head, midpiece and proximal portion of flagellum) of Hemidonax pictus (Tryon, 1870) (A), Cardiidae (BK), Donacidae (LP) and selected Veneridae (QV). BK, Cardiidae. B, Fragum unedo (Linnaeus, 1758); C, Lunulicardia hemicardium (Linnaeus, 1758); D, Vasticardium vertebratum (Jonas, 1844); E, Papyridea semisulcata (Gray, 1825); F, Fulvia tenuicostata (Lamarck, 1819); G, H, Cerastoderma spp.; I, Hippopus hippopus (Linnaeus, 1758); J, Tridacna gigas (Linnaeus, 1758); K, Tridacna maxima (Röding, 1798). LP, Donacidae. L, Donax deltoides Lamarck, 1818; M, Donax madagascariensis Wood, 1828; N, Donax sordidus Hanley, 1845; O, Donax serra Röding, 1798; P, Donax trunculus Linnaeus, 1758. QV, Veneridae. Q, Venerupis aurea (Gmelin, 1791); R, Venerupis corrugata (Gmelin, 1791); S, Petricola lapicida (Gmelin, 1791); T, Antigona chemnitzii (Hanley, 1844); U, Dosinia nedigna (Iredale, 1930); V, Lioconcha annettae Lamprell &amp; Whitehead, 1990. Scale bar for Hemidonax pictus (= 1.0 Mm) applies to all other figures except H (= 2.0 Mm). Sources of data: AE, L, SV (this paper; J. M. Healy, P. M. Mikkelsen &amp; R. Bieler, unpubl. data); F (Popham, 1979); G, H [composite reconstruction based on Cerastoderma lamarcki (Reeve, 1844) from Drozdov et al., 2001 and Cerastoderma edule (Linnaeus, 1758) from Sousa &amp; Azevedo, 1988 and Sousa et al., 1998]; IK (Keys &amp; Healy, 1999, 2000); MO (Hodgson et al., 1990); P (Sousa &amp; Oliveira, 1994); Q, R (Pochon-Masson &amp; Gharagozlou, 1970; Gharagozlou-Van Ginneken &amp; Pochon-Masson, 1971)." pageId="10" pageNumber="335">Fig. 8</figureCitation>
, also for comparative figures and extensive literature see
<bibRefCitation id="FF204B05633F1E2D1FE74ECAFC65F96D" author="Healy JM" box="[1161,1318,1695,1717]" pageId="10" pageNumber="335" pagination="155 - 166" refId="ref11712" refString="Healy JM. 1995 b. Comparative sperm ultrastructure in veneroid bivalves (Mollusca). In: Jamieson BGM, Ausio J, Justine J-L, eds. Advances in spermatozoal phylogeny and taxonomy. Memoirs du Museum National d'Histoire Naturelle 166: 155 - 166." type="journal article" year="1995">Healy, 1995b</bibRefCitation>
,
<bibRefCitation id="FF204B05633F1E2D1E544ECAFCC2F96D" author="Healy JM" box="[1338,1409,1695,1717]" pageId="10" pageNumber="335" pagination="99 - 113" refId="ref11760" refString="Healy JM. 1996 a. Molluscan sperm ultrastructure: correlation with taxonomic units within the Gastropoda, Cephalopoda and Bivalvia. In: Taylor J, ed. Origin and evolutionary radiation of the Mollusca. Oxford: Oxford University Press, 99 - 113." type="book chapter" year="1996">1996a</bibRefCitation>
). Substantial diversity exists among heterodont taxa in the shape of the acrosomal vesicle (and the shape and internal structure of the basal ring), as well as the size and length of the nucleus (short or long, straight, curved or helical) and the number of midpiece mitochondria (four or five, sometimes more, but always with a predominating number). As there are a number of differing opinions relating to the placement and affinities of
<taxonomicName id="5CB14D77633E1E2C19E64F6FFA49F897" authorityName="Morch" authorityYear="1778" box="[648,778,1850,1871]" class="Bivalvia" family="Hemidonacidae" genus="Hemidonax" kingdom="Animalia" order="Venerida" pageId="11" pageNumber="336" phylum="Mollusca" rank="genus">
<emphasis id="A9C5EAE6633E1E2C19E64F6FFA49F897" box="[648,778,1850,1871]" italics="true" pageId="11" pageNumber="336">Hemidonax</emphasis>
</taxonomicName>
among the Heterodonta, the following part of the discussion will deal with each of these, including a brief resumé of group features then progressing to a comparison with
<taxonomicName id="5CB14D77633E1E2C1E714F6FFCE2F897" authorityName="Morch" authorityYear="1778" box="[1311,1441,1850,1871]" class="Bivalvia" family="Hemidonacidae" genus="Hemidonax" kingdom="Animalia" order="Venerida" pageId="11" pageNumber="336" phylum="Mollusca" rank="genus">
<emphasis id="A9C5EAE6633E1E2C1E714F6FFCE2F897" box="[1311,1441,1850,1871]" italics="true" pageId="11" pageNumber="336">Hemidonax</emphasis>
</taxonomicName>
results.
</paragraph>
</subSubSection>
</treatment>
</document>