<mods:titleid="945FFEC11C12CB37C7896501F1FD962B">Taxonomic review of the Spixia pyriformis species complex (Gastropoda: Pulmonata: Odontostominae)</mods:title>
<bibRefCitationid="B54D01F73B057E2F5276FB53FDCBFA83"author="Doering"box="[364,534,1271,1297]"pageId="8"pageNumber="25"refString="Doering, A. (1874). Apuntes sobre la fauna de moluscos de la Republica Argentina. Boletin de Academia Nacional de Ciencias, Cordoba, 1, 452 - 457."type="journal article"year="1874">Doering 1874</bibRefCitation>
<figureCitationid="49E760833B057E2F5385FABEFE97FAA6"box="[159,330,1306,1332]"captionStart-0="FIGURES 15 – 20"captionStart-1="FIGURES 21 – 23"captionStart-2="FIGURE 24"captionStart-3="FIGURES 25 – 26"captionStartId-0="9.[151,269,1717,1741]"captionStartId-1="11.[151,269,1826,1850]"captionStartId-2="12.[151,255,1988,2012]"captionStartId-3="13.[151,269,1615,1639]"captionTargetBox-0="[264,1323,514,1693]"captionTargetBox-1="[292,1300,345,1814]"captionTargetBox-2="[262,1324,554,1964]"captionTargetBox-3="[262,1324,184,1591]"captionTargetId-0="figure@9.[264,1323,514,1693]"captionTargetId-1="figure@11.[292,1301,332,1815]"captionTargetId-2="figure@12.[262,1325,554,1964]"captionTargetId-3="figure@13.[262,1325,184,1591]"captionTargetPageId-0="9"captionTargetPageId-1="11"captionTargetPageId-2="12"captionTargetPageId-3="13"captionText-0="FIGURES 15 – 20. Spixia martensii. 15. Paratype MACN 23139.16. Dorsal view of protoconch and next whorl smooth, scale bar: 100 Μm. 17. View from protoconch to sixth whorl of a juvenile shell. Note spiral rows and lamellae, scale bar: 1 mm. 18. Third and fourth whorl with spiral major rows with lamellae and minor files, scale bar: 1 mm. 19. Detail of sculpture in fourth whorl, spiral major row with lamella and minor rows with micro folds, arrow points to horizontal fine line, scale bar: 100 Μm. 20. Jaw with central plaques, scale bar: 100 Μm."captionText-1="FIGURES 21 – 23. Spixia martensii. 21. Reproductive system dissected out, scale bar: 5 mm. 22. Phallic complex inner wall, scale bar: 5 mm. 23. Detail of constriction and widening area II and part of area III, scale bar: 2 mm. Abbreviations: A: atrium; Ag: albumen gland; Bc: bursa copulatrix; Bcd: bursa copulatrix duct; Ca: constriction area ;; Ep: epiphallus; F: flagellum; Hd: hermaphroditic duct; Lf: longitudinal folds; Ov: oviduct; P: penis; Pg: penial gland; Pi: pilaster; Piw: penis inner wall; Pr: prostate; Pro: projections; Ps: penial sheath; Piw: penis inner wall; Rm: retractor muscle I: proximal portion; II: widening area; III: proximal medial portion; IV: distal medial portion; V: distal portion."captionText-2="FIGURE 24. Spixia martensii: Size and shape variation of shells, scale bar: 10 mm."captionText-3="FIGURES 25 – 26. Spixia martensii. 25. Shell with and without periostracum, arrow points to spiral row, scale bar: 1 mm. 26. Detail of shell with and without periostracum showing the grooves left by spiral rows, arrow points to groove, scale bar: 1 mm."httpUri-0="https://zenodo.org/record/177065/files/figure.png"httpUri-1="https://zenodo.org/record/177066/files/figure.png"httpUri-2="https://zenodo.org/record/177067/files/figure.png"httpUri-3="https://zenodo.org/record/177068/files/figure.png"pageId="8"pageNumber="9">Figures 15–26</figureCitation>
<bibRefCitationid="B54D01F73B057E2F52B9FACEFD9DFA13"author="Doering"box="[419,576,1386,1409]"pageId="8"pageNumber="25"refString="Doering, A. (1874). Apuntes sobre la fauna de moluscos de la Republica Argentina. Boletin de Academia Nacional de Ciencias, Cordoba, 1, 452 - 457."type="journal article"year="1874">Doering, 1874</bibRefCitation>
<bibRefCitationid="B54D01F73B057E2F52B9FA2EFD9DFA33"author="Doering"box="[419,576,1418,1441]"pageId="8"pageNumber="25"refString="Doering, A. (1875) Estudios Sistematicos y Anatomicos sobre Moluscos Pulmoniferos de Los Paises del Plata. Periodico Zoologico, 3, 171 - 198."type="journal article"year="1875">Doering, 1875</bibRefCitation>
<bibRefCitationid="B54D01F73B057E2F5109FA6EFD74FA73"author="Pfeiffer"box="[531,681,1482,1505]"pageId="8"pageNumber="25"refString="Pfeiffer, L. (1877) Monographia Helicorium viventum. Sistens desriptiones systematicas et criticas omnium huius familiae generum et specierum hodie cognitarum. Lipsiae (Brockhaus), 8, 10 + 674 pp."type="book"year="1877">Pfeiffer, 1877</bibRefCitation>
<bibRefCitationid="B54D01F73B057E2F52E4FA4EFD51F993"author="Pilsbry"box="[510,652,1514,1537]"pageId="8"pageNumber="25"refString="Pilsbry, H. A. (1901) Manual of Conchology, 2 ª serie, Pulmonata, Philadelphia, XIV, pp. 1 - 320."type="journal article"year="1901">Pilsbry, 1901</bibRefCitation>
<bibRefCitationid="B54D01F73B057E2F52E4F9AEFD7EF9B3"author="Parodiz"box="[510,675,1546,1569]"pageId="8"pageNumber="25"refString="Parodiz, J. J. (1942 b) Los Odontostominos en la Argentina, Segunda parte, Physis, 19, 319 - 343."type="journal article"year="1942"yearSuffix="b">Parodiz, 1942b</bibRefCitation>
<bibRefCitationid="B54D01F73B057E2F52EEF98EFD56F9D3"author="Parodiz"box="[500,651,1578,1601]"pageId="8"pageNumber="25"refString="Parodiz, J. J. (1957) Catalogue of the land Mollusca of Argentina. The Nautilus, 71, 22 - 31."type="journal article"year="1957">Parodiz, 1957</bibRefCitation>
<bibRefCitationid="B54D01F73B057E2F52FCF9EEFDB8F9F3"author="Zilch"box="[486,613,1610,1633]"pageId="8"pageNumber="25"refString="Zilch, A. (1971) Die Typen und Typoide des Natur-Museums Seckenberg, 47: Mollusca Euthyneura von A. Doring aus Argentinien. Archiv fur Molluskenkunde, 101, 195 - 213."type="journal article"year="1971">Zilch, 1971</bibRefCitation>
<bibRefCitationid="B54D01F73B057E2F5256F9CEFDDCF913"author="Fernandez"box="[332,513,1642,1665]"pageId="8"pageNumber="25"refString="Fernandez, D. (1973) Catalogo de la Malacofauna Terrestre Argentina. Comision de Investigaciones Cientificas de la Provincia de Buenos Aires. Provincia de Buenos Aires Press, La Plata, Argentina, 197 pp."type="book"year="1973">Fernández, 1973</bibRefCitation>
<figureCitationid="49E760833B057E2F520BF88EFEB6F8D6"box="[273,363,1834,1860]"captionStart="FIGURES 15 – 20"captionStartId="9.[151,269,1717,1741]"captionTargetBox="[264,1323,514,1693]"captionTargetId="figure@9.[264,1323,514,1693]"captionTargetPageId="9"captionText="FIGURES 15 – 20. Spixia martensii. 15. Paratype MACN 23139.16. Dorsal view of protoconch and next whorl smooth, scale bar: 100 Μm. 17. View from protoconch to sixth whorl of a juvenile shell. Note spiral rows and lamellae, scale bar: 1 mm. 18. Third and fourth whorl with spiral major rows with lamellae and minor files, scale bar: 1 mm. 19. Detail of sculpture in fourth whorl, spiral major row with lamella and minor rows with micro folds, arrow points to horizontal fine line, scale bar: 100 Μm. 20. Jaw with central plaques, scale bar: 100 Μm."httpUri="https://zenodo.org/record/177065/files/figure.png"pageId="8"pageNumber="9">Fig. 15</figureCitation>
): Subpyriform elongated, 8 1/2 whorls, first ones convex, less convexity in last two ones. Color light brown slightly cupreous with irregular whitish axial bands. Protoconch with slim, axial ribs, parallel, slightly curved on superior edge, among ribs, ultraestructure surface consisting in perpendicularly micro bands cutting axial ribs (
<figureCitationid="49E760833B057E2F52A4F806FDCAF82E"box="[446,535,1954,1980]"captionStart="FIGURES 15 – 20"captionStartId="9.[151,269,1717,1741]"captionTargetBox="[264,1323,514,1693]"captionTargetId="figure@9.[264,1323,514,1693]"captionTargetPageId="9"captionText="FIGURES 15 – 20. Spixia martensii. 15. Paratype MACN 23139.16. Dorsal view of protoconch and next whorl smooth, scale bar: 100 Μm. 17. View from protoconch to sixth whorl of a juvenile shell. Note spiral rows and lamellae, scale bar: 1 mm. 18. Third and fourth whorl with spiral major rows with lamellae and minor files, scale bar: 1 mm. 19. Detail of sculpture in fourth whorl, spiral major row with lamella and minor rows with micro folds, arrow points to horizontal fine line, scale bar: 100 Μm. 20. Jaw with central plaques, scale bar: 100 Μm."httpUri="https://zenodo.org/record/177065/files/figure.png"pageId="8"pageNumber="9">Fig. 16</figureCitation>
). Second whorl from protoconch, smooth in appearance. Teleoconch sculpture with axial oblique wrinkles. From third whorl to sixth, three spiral major rows, elevated, not continuous, on each whorl. Triangular lamellae inserted on these major rows (
<figureCitationid="49E760833B047E2E5082FF3CFBC9FF20"box="[920,1044,152,178]"captionStart="FIGURES 15 – 20"captionStartId="9.[151,269,1717,1741]"captionTargetBox="[264,1323,514,1693]"captionTargetId="figure@9.[264,1323,514,1693]"captionTargetPageId="9"captionText="FIGURES 15 – 20. Spixia martensii. 15. Paratype MACN 23139.16. Dorsal view of protoconch and next whorl smooth, scale bar: 100 Μm. 17. View from protoconch to sixth whorl of a juvenile shell. Note spiral rows and lamellae, scale bar: 1 mm. 18. Third and fourth whorl with spiral major rows with lamellae and minor files, scale bar: 1 mm. 19. Detail of sculpture in fourth whorl, spiral major row with lamella and minor rows with micro folds, arrow points to horizontal fine line, scale bar: 100 Μm. 20. Jaw with central plaques, scale bar: 100 Μm."httpUri="https://zenodo.org/record/177065/files/figure.png"pageId="9"pageNumber="10">Fig.17–18</figureCitation>
). Between each major row, 4 to 6 spiral minor rows slightly undulated in zigzag, separated by regular spaces. Departing from each row, grow axial micro folds, not straight, connecting each other in the center; under them horizontal fine lines form a grillwork together with the previous ones (
<figureCitationid="49E760833B047E2E5190FEB4FD3DFEB8"box="[650,736,272,298]"captionStart="FIGURES 15 – 20"captionStartId="9.[151,269,1717,1741]"captionTargetBox="[264,1323,514,1693]"captionTargetId="figure@9.[264,1323,514,1693]"captionTargetPageId="9"captionText="FIGURES 15 – 20. Spixia martensii. 15. Paratype MACN 23139.16. Dorsal view of protoconch and next whorl smooth, scale bar: 100 Μm. 17. View from protoconch to sixth whorl of a juvenile shell. Note spiral rows and lamellae, scale bar: 1 mm. 18. Third and fourth whorl with spiral major rows with lamellae and minor files, scale bar: 1 mm. 19. Detail of sculpture in fourth whorl, spiral major row with lamella and minor rows with micro folds, arrow points to horizontal fine line, scale bar: 100 Μm. 20. Jaw with central plaques, scale bar: 100 Μm."httpUri="https://zenodo.org/record/177065/files/figure.png"pageId="9"pageNumber="10">Fig. 19</figureCitation>
). Suture simple, slightly pronounced. Aperture ovate, wide, peristome slightly reflex. Parietal callus present, of medium development. Aperture with one tooth, four lamellae. Suprapalatal tooth shallowly developed or absent. Palatal lamellae with concave sides and bulky edge, slightly oblique respect peristome. Basal lamellae narrow. Columellar lamellae concave in its superior side, perpendicular, oblique from peristome. Parietal lamellae stretched (
<figureCitationid="49E760833B047E2E50F0FE14FB9CFE58"box="[1002,1089,432,458]"captionStart="FIGURES 15 – 20"captionStartId="9.[151,269,1717,1741]"captionTargetBox="[264,1323,514,1693]"captionTargetId="figure@9.[264,1323,514,1693]"captionTargetPageId="9"captionText="FIGURES 15 – 20. Spixia martensii. 15. Paratype MACN 23139.16. Dorsal view of protoconch and next whorl smooth, scale bar: 100 Μm. 17. View from protoconch to sixth whorl of a juvenile shell. Note spiral rows and lamellae, scale bar: 1 mm. 18. Third and fourth whorl with spiral major rows with lamellae and minor files, scale bar: 1 mm. 19. Detail of sculpture in fourth whorl, spiral major row with lamella and minor rows with micro folds, arrow points to horizontal fine line, scale bar: 100 Μm. 20. Jaw with central plaques, scale bar: 100 Μm."httpUri="https://zenodo.org/record/177065/files/figure.png"pageId="9"pageNumber="10">Fig. 15</figureCitation>
. 15. Paratype MACN 23139.16. Dorsal view of protoconch and next whorl smooth, scale bar: 100 Μm. 17. View from protoconch to sixth whorl of a juvenile shell. Note spiral rows and lamellae, scale bar: 1 mm. 18. Third and fourth whorl with spiral major rows with lamellae and minor files, scale bar: 1 mm. 19. Detail of sculpture in fourth whorl, spiral major row with lamella and minor rows with micro folds, arrow points to horizontal fine line, scale bar: 100 Μm. 20. Jaw with central plaques, scale bar: 100 Μm.
: Jaw arcuate, light brown, lustrous, formed by 16 imbricate plaques (
<figureCitationid="49E760833B077E2D57CBFE2CFAF7FE30"box="[1233,1322,392,418]"captionStart="FIGURES 15 – 20"captionStartId="9.[151,269,1717,1741]"captionTargetBox="[264,1323,514,1693]"captionTargetId="figure@9.[264,1323,514,1693]"captionTargetPageId="9"captionText="FIGURES 15 – 20. Spixia martensii. 15. Paratype MACN 23139.16. Dorsal view of protoconch and next whorl smooth, scale bar: 100 Μm. 17. View from protoconch to sixth whorl of a juvenile shell. Note spiral rows and lamellae, scale bar: 1 mm. 18. Third and fourth whorl with spiral major rows with lamellae and minor files, scale bar: 1 mm. 19. Detail of sculpture in fourth whorl, spiral major row with lamella and minor rows with micro folds, arrow points to horizontal fine line, scale bar: 100 Μm. 20. Jaw with central plaques, scale bar: 100 Μm."httpUri="https://zenodo.org/record/177065/files/figure.png"pageId="10"pageNumber="11">Fig. 20</figureCitation>
). Plaques show fine transversal irregular lines. Most external plaques double size respect to central ones. Inferior border of jaw with irregular outline due to projections of plaques. Radula identical with
<figureCitationid="49E760833B077E2D52C8FDF4FDFAFDF8"box="[466,551,592,618]"captionStart="FIGURES 21 – 23"captionStartId="11.[151,269,1826,1850]"captionTargetBox="[292,1300,345,1814]"captionTargetId="figure@11.[292,1301,332,1815]"captionTargetPageId="11"captionText="FIGURES 21 – 23. Spixia martensii. 21. Reproductive system dissected out, scale bar: 5 mm. 22. Phallic complex inner wall, scale bar: 5 mm. 23. Detail of constriction and widening area II and part of area III, scale bar: 2 mm. Abbreviations: A: atrium; Ag: albumen gland; Bc: bursa copulatrix; Bcd: bursa copulatrix duct; Ca: constriction area ;; Ep: epiphallus; F: flagellum; Hd: hermaphroditic duct; Lf: longitudinal folds; Ov: oviduct; P: penis; Pg: penial gland; Pi: pilaster; Piw: penis inner wall; Pr: prostate; Pro: projections; Ps: penial sheath; Piw: penis inner wall; Rm: retractor muscle I: proximal portion; II: widening area; III: proximal medial portion; IV: distal medial portion; V: distal portion."httpUri="https://zenodo.org/record/177066/files/figure.png"pageId="10"pageNumber="11">Fig. 21</figureCitation>
) Ovotestis consisting of five bunches of acini. Each one fan shaped, digitiform, ramified, orange-yellowish homogeneous. Free oviduct short and narrow 1/6 of spermoviduct total length. Bursa copulatrix duct 1/4 longer than spermoviduct, with distal portion enlarged. Vagina 1/3 of penis length, inner wall consisting of folds with small irregular pustules.
Phallic complex comprised by penis, epiphallus, and flagellum. Flagellum short, filiform. Epiphallus twice the length and diameter of flagellum. Inner wall of penial area I with small folds (
<figureCitationid="49E760833B077E2D57ADFCBCFAE0FCA0"box="[1207,1341,792,818]"captionStart="FIGURES 21 – 23"captionStartId="11.[151,269,1826,1850]"captionTargetBox="[292,1300,345,1814]"captionTargetId="figure@11.[292,1301,332,1815]"captionTargetPageId="11"captionText="FIGURES 21 – 23. Spixia martensii. 21. Reproductive system dissected out, scale bar: 5 mm. 22. Phallic complex inner wall, scale bar: 5 mm. 23. Detail of constriction and widening area II and part of area III, scale bar: 2 mm. Abbreviations: A: atrium; Ag: albumen gland; Bc: bursa copulatrix; Bcd: bursa copulatrix duct; Ca: constriction area ;; Ep: epiphallus; F: flagellum; Hd: hermaphroditic duct; Lf: longitudinal folds; Ov: oviduct; P: penis; Pg: penial gland; Pi: pilaster; Piw: penis inner wall; Pr: prostate; Pro: projections; Ps: penial sheath; Piw: penis inner wall; Rm: retractor muscle I: proximal portion; II: widening area; III: proximal medial portion; IV: distal medial portion; V: distal portion."httpUri="https://zenodo.org/record/177066/files/figure.png"pageId="10"pageNumber="11">Fig. 22–23</figureCitation>
). Penial gland short, rounded. Penial area II with two ovate projections of numerous festoon shaped rows circularly arranged leaving circular small spaces. Within distal portion of area II a lateral pilaster running towards to area IV is present. Wall of penial area III with rows in zigzag, V shaped, becomes longitudinal towards area V. Penial area V smooth, with longitudinal folds, semi undulated, until atrium. Retractor muscle inserted externally at proximal portion of area II. Vas deferens short, stacked to retractor muscle; running independently parallel to epiphallus. Penial sheath short thin, located at distal portion of penis.
<bibRefCitationid="B54D01F73B077E2D574DFB24FACAFB08"author="Morrone"box="[1111,1303,1152,1178]"pageId="10"pageNumber="25"refString="Morrone, J. J. (2006) Biogeografic Areas And Transitions Zones of Latin America And The Caribean Islands Based On Pambiogeografic And Cladistic Analyses of The Entomofauna. Annual Review of Entomology, 51, 467 - 494."type="journal article"year="2006">Morrone (2006)</bibRefCitation>
is highly variable in shell size. Due to the variability in the body whorl’s diameter within the present species, the subpyriform shell shape may not be so evident in some specimens (
<figureCitationid="49E760833B077E2D5385FB5CFF2AFA80"box="[159,247,1272,1298]"captionStart="FIGURE 24"captionStartId="12.[151,255,1988,2012]"captionTargetBox="[262,1324,554,1964]"captionTargetId="figure@12.[262,1325,554,1964]"captionTargetPageId="12"captionText="FIGURE 24. Spixia martensii: Size and shape variation of shells, scale bar: 10 mm."httpUri="https://zenodo.org/record/177067/files/figure.png"pageId="10"pageNumber="11">Fig. 24</figureCitation>
). Shell wrinkles, that are characteristics of the body whorl in this species, are sometimes not conspicuous even when the periostracum is well preserved. Shell sculpture until 6 1/2 whorl on both adults and juveniles, is similar to that of
collected alive, a marked change of shell sculpture at the level of the sixth whorl was observed. The spiral rows are replaced by grooves that combined with axial wrinkles give a network appearance to the last whorls surface. In some specimens, although having a complete periostracum, the spiral rows disappear at whorl six with vestiges of spiral grooves left (
<figureCitationid="49E760833B077E2D5157F99CFD08F9C0"box="[589,725,1592,1618]"captionStart="FIGURES 25 – 26"captionStartId="13.[151,269,1615,1639]"captionTargetBox="[262,1324,184,1591]"captionTargetId="figure@13.[262,1325,184,1591]"captionTargetPageId="13"captionText="FIGURES 25 – 26. Spixia martensii. 25. Shell with and without periostracum, arrow points to spiral row, scale bar: 1 mm. 26. Detail of shell with and without periostracum showing the grooves left by spiral rows, arrow points to groove, scale bar: 1 mm."httpUri="https://zenodo.org/record/177068/files/figure.png"pageId="10"pageNumber="11">Fig. 25–26</figureCitation>
was not previously described or illustrated before by any author and the spiral rows and lamellae inserted on them where not mentioned either in previous descriptions. The pallial gland is usually smaller than in
and different from the rest of the species complex where the penial gland is short and triangular. The inner sculpture of penial wall is different from the other species except for penial areas I and V, that are similar to the other species. The penial area II is completely different from the other species, because the present species has a different arrangement of the rows, ordered into two conspicuous projections festoon shaped. The penial area III and area IV in
but showing V-shaped rows more twisted and tighter. In the present species the insertion point of retractor muscle is at proximal portion of penial area II, similar than in
locality in Doering´s original description is not clearly specified. He indicated it as “Cordoba, Totoral”. However, Totoral is an entire political department of Cordoba Province in central
. In Totoral department neither dead nor alive material was found by the author. Also, most of the material currently deposited at the Museums, except for the
, is from Punilla, Ischilín, Tulumba and Calamuchita departments from Cordoba.
<bibRefCitationid="B54D01F73B017E2B52F3FE9CFD74FEC0"author="Parodiz"box="[489,681,312,338]"pageId="12"pageNumber="25"refString="Parodiz, J. J. (1942 b) Los Odontostominos en la Argentina, Segunda parte, Physis, 19, 319 - 343."type="journal article"year="1942"yearSuffix="b">Parodiz (1942b)</bibRefCitation>
ranges from Cordoba, Catamarca to Tucumán. Nevertheless, no material of this species was yet found alive in field in Catamarca and Tucumán provinces. We now can extend the range of this species to Salta Province in north-western
, where a group of specimens was found close to Juramento River. New field work and extensive collections are needed to precise the real distribution of
. 25. Shell with and without periostracum, arrow points to spiral row, scale bar: 1 mm. 26. Detail of shell with and without periostracum showing the grooves left by spiral rows, arrow points to groove, scale bar: 1 mm.
