Abstract
Santaremicaris gen. nov. is proposed to accommodate Santaremicaris amazonensis sp. nov., a new species collected by Noodt from the hyporheic zone of the Amazonas River at Santarém, Brazil, and Parastenocaris santaremensis Noodt, 1963, which is redescribed and designated as the type species of the new genus. The new genus is recorded also from several localities of the Amazonian hydrographic basin. Santaremicaris gen. nov. is well supported by clear autapomorphies, e.g., 1) the absence of distal spinules on the endopod of the second swimming leg of the male, which is ornamented with strong outer spinules along the first 2/3 of the segment and armed with a long distal seta the insertion area of which covers almost the whole width of the segment; 2) the presence of a triangular, short and strongly sclerotized endopod on the fourth swimming leg of the male; and 3) the presence of a long and slim male fifth leg ending in a long distal spiniform process, with irregular margins, armed with only three setigerous elements on the outer margin, and reaching the middle of the subsequent urosomal segment or beyond it. The new genus is closely related to Murunducaris Reid, 1994, with males sharing conspicuous ornamentation on the inner margin of the first swimming leg basis, represented either by an enlarged spinule accompanied by minute accessory spinules or by a row of strong spinules. In addition, the males of both Murunducaris and Santaremicaris gen. nov. share an enlarged intercoxal sclerite with a median process on the fifth leg.
Keywords
Amazonian basin; Fontinalicaridinae; groundwater; hyporheos; Neotropics
INTRODUCTION
Knowledge of taxonomy and phylogenetic affinities within the family Parastenocarididae is advancing in recent years. Several new genera were described (i.e., SimplicarisGalassi and De Laurentiis, 2004Galassi, D.M.P. and de Laurentiis, P. 2004. Toward a revision of the genus Parastenocaris Kessler, 1913: estabilishment of Simplicaris gen. nov. from groundwaters in central Italy and review of the P. brevipes-group (Copepoda, Harpacticoida, Parastenocarididae). Zoological Journal of the Linean Society, 140: 417-436.; MonodicarisSchminke, 2009Schminke, H.K. 2009. Monodicaris gen. n. (Copepoda, Harpacticoida, Parastenocarididae) from west Africa. Crustaceana, 82: 367-378; AsiacarisCottarelli, Bruno and Berera, 2010Cottarelli, V.; Bruno, M.C. and Berera, R. 2010. First record of Parastenocarididae from Thailand and description of a new genus (Copepoda: Harpacticoida). Journal of Crustacean Biology, 30: 478-494.; DussartstenocarisKaranovic and Cooper, 2011Karanovic, T. and Cooper, S.J.B. 2011. Third genus of parastenocaridid copepods from Australia supported by molecular evidence (Copepoda, Harpacticoida). In: Defaye D.; Suárez-Morales, E. and Vaupel Klein J.C. (eds), Studies on Freshwater Copepoda: a Volume in Honour of Bernard Dussart Crustaceana Monographs, 305-321. ; IticocarisCorgosinho, Martínez Arbizu and Previatelli, 2012aCorgosinho, P.H.C.; Martínez Arbizu, P. and Previatelli, D. 2012a. Establishment of a new genus for Parastenocaris itica (Copepoda, Harpacticoida) from El Salvador, Central America, with discussion of the Parastenocaris fontinalis and P. proserpina groups. Iheringia, Série Zoologia, 102: 401-411. ; Horstkurtcaris Karanovic and Lee, 2012Karanovic, T. and Lee, W. 2012. A new species of Parastenocaris from Korea, with a redescription of the closely related P. biwae from Japan (Copepoda: Harpacticoida: Parastenocarididae). Journal of Species Research, 1: 4-34.; CottarellicarisSchminke, 2013Schminke, H.K. 2013. Stammericaris Jakobi, 1972 redefined and a new genus of Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 86: 704-717.; HimalayacarisRanga Reddy, Totakura and Corgosinho, 2014Totakura, V.R.; Ranga-Reddy, Y. and Shaik, S. 2014. Three new species of Parastenocarididae (Crustacea, Copepoda, Harpacticoida) from India. Zootaxa, 3821: 501-537.; IndocarisRanga Reddy, Totakura and Shaik, 2016Ranga Reddy, Y.; Totakura, V.R. and Shaik, S. 2016. A new genus and two new species of Parastenocarididae (Copepoda: Harpacticoida) from southeastern India. Journal of Natural History, 50: 1315-1356.; EirinicarisCorgosinho, Schizas, Previattelli, Rocha and Santos-Silva, 2017bCorgosinho, P.H.C.; Schizas, N.V.; Previattelli, D.; Rocha, C.E.F. da and Santos-Silva, E.N. 2017b. A new genus of Parastenocarididae (Copepoda, Harpacticoida) from the Tocantins River basin (Goiás, Brazil), and a phylogenetic analysis of the Parastenocaridinae. Zoosystematics and Evolution, 93: 167-187.; ColombocarisGaviria, Defaye and Corgosinho, 2017Gaviria, S.; Defaye, D. and Corgosinho, P.H.C. 2017. Two new parastenocaridid genera (Copepoda, Harpacticoida) from South America, including a revision of Noodt’s columbiensis-group and the redescription of Parastenocaris tapajoensis Noodt, 1963. Crustaceana, 90: 535-570. ; and NoodtcarisGaviria, Defaye and Corgosinho, 2017Gaviria, S.; Defaye, D. and Corgosinho, P.H.C. 2017. Two new parastenocaridid genera (Copepoda, Harpacticoida) from South America, including a revision of Noodt’s columbiensis-group and the redescription of Parastenocaris tapajoensis Noodt, 1963. Crustaceana, 90: 535-570. ). The classic work of Jakobi (1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146.) was revisited and many genera were redefined and reestablished [i.e., RemaneicarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146. (Corgosinho and Martínez Arbizu, 2005Corgosinho, P.H.C. and Martínez Arbizu, P. 2005. Two new interstitial species of Remaneicaris Jakobi (Copepoda, Harpacticoida, Parastenocarididae) from the Ribeirão do Ouro river, Brazil, with a redefinition of the genus. Senckenbergiana biologica, 85: 147-162.); BrasilibathynellocarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146. (Corgosinho et al., 2010aCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2010a. Revision of Brasilibathynellocaris Jakobi, 1972 (Copepoda: Harpacticoida: Parastenocarididae) with redefinition of the genus. Zoological Journal of the Linnean Society, 159: 527-566.); KinnecarisJakobi, 1972 Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146.(Schminke, 2008Schminke, H.K. 2008. First report of groundwater fauna from Papua New Guinea: Kinnecaris Jakobi, 1972 redefined (Copepoda, Harpacticoida, Parastenocarididae), and description of a new species. Crustaceana, 81: 1241-1253.); SiolicarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146. (Corgosinho et al., 2012bCorgosinho, P.H.C.; Ranga Reddy, Y. and Martínez Arbizu, P. 2012b. Revision of the genus Siolicaris Jakobi, 1972, with redescriptions of S. sioli (Noodt, 1963) and S. jakobi (Noodt, 1963) from South America, and S. sandhya (Ranga Reddy, 2001) comb. nov. from India (Copepoda, Harpacticoida, Parastenocarididae). Zootaxa, 3493: 49-71.); ProserpinicarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146. (Karanovic, Cho and Lee, 2012Karanovic, T.; Cho, J-L. and Lee, W. 2012. Redefinition of the parastenocaridid genus Proserpinicaris (Copepoda: Harpacticoida), with description of three new species from Korea. Journal of Natural History, 46: 25-26.); StammericarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146. (Schminke, 2013Schminke, H.K. 2013. Stammericaris Jakobi, 1972 redefined and a new genus of Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 86: 704-717.), ParastenocarisKessler, 1913Kessler, E. 1913. Parastenocaris brevipes nov. gen. et nov. spec., ein neuer Süsswasserharpacticide. Zoologischer Anzeiger, 42(11): 514-520. (Karanovic and Lee, 2012Karanovic, T. and Lee, W. 2012. A new species of Parastenocaris from Korea, with a redescription of the closely related P. biwae from Japan (Copepoda: Harpacticoida: Parastenocarididae). Journal of Species Research, 1: 4-34.)]. Consequently, PararemaneicarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146. and ParaforficatocarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146. were synonymized with Brasilibathynellocaris (see Corgosinho et al., 2010aCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2010a. Revision of Brasilibathynellocaris Jakobi, 1972 (Copepoda: Harpacticoida: Parastenocarididae) with redefinition of the genus. Zoological Journal of the Linnean Society, 159: 527-566.); NipponicarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146. and PannonicarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146. were synonymized with Proserpinicaris (see Karanovic, Cho and Lee, 2012Karanovic, T.; Cho, J-L. and Lee, W. 2012. Redefinition of the parastenocaridid genus Proserpinicaris (Copepoda: Harpacticoida), with description of three new species from Korea. Journal of Natural History, 46: 25-26.); BiwaecarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146., OshimaensicarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146., EnckellicarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146., and BrinckicarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146. were synonymized with Parastenocaris (see Karanovic and Lee, 2012Karanovic, T. and Lee, W. 2012. A new species of Parastenocaris from Korea, with a redescription of the closely related P. biwae from Japan (Copepoda: Harpacticoida: Parastenocarididae). Journal of Species Research, 1: 4-34.); CafferocarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146. was synonymized with Kinnecaris (see Schminke, 2013Schminke, H.K. 2013. Stammericaris Jakobi, 1972 redefined and a new genus of Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 86: 704-717.); and PhreaticarisJakobi, 1972Jakobi, H. 1972. Trends (Enp. P4 ♂) innerhalb der Parastenocarididen (Copepoda, Harpacticoida). Crustaceana, 22: 127-146. was synonymized with Stammericaris (see Schminke, 2013Schminke, H.K. 2013. Stammericaris Jakobi, 1972 redefined and a new genus of Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 86: 704-717.). Despite the work of Karanovic and Lee (2012Karanovic, T. and Lee, W. 2012. A new species of Parastenocaris from Korea, with a redescription of the closely related P. biwae from Japan (Copepoda: Harpacticoida: Parastenocarididae). Journal of Species Research, 1: 4-34.) in redefining the genus Parastenocaris, with the exception of the species belonging to the Parastenocaris brevipes-group, the genus Parastenocaris still remains as a repository of loosely related species of unclear phylogenetic affinities within the subfamilies Fontinalicaridinae Schminke, 2010 Schminke, H.K. 2010. High-level phylogenetic relationships within Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 83: 343-367.and Parastenocaridinae Chappuis, 1940Chappuis, P.A. 1940. Die Harpacticoiden des Grundwassers des unteren Maintales. Archiv für Hidrobiologie, 36: 286-305. (see Schminke, 2010Schminke, H.K. 2010. High-level phylogenetic relationships within Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 83: 343-367.).
The Neotropical fauna of Parastenocarididae is composed of about 74 described species and 11 genera (Walter and Boxshall, 2020Walter, T.C. and Boxshall, G. 2020. World of Copepods database. Parastenocarididae Chappuis, 1940. Accessed through: World Register of Marine Species at: Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=taxdetails&id=115170 . Acessed on 05 november 2020.
http://www.marinespecies.org/aphia.php?p...
). PotamocarisDussart, 1979Dussart, B.H. 1979. Algunos copepodos de America del Sur. Museo Nacional de Historia Natural, Santiago de Chile, Publicación Ocasional, 30: 3-13., ForficatocarisJakobi, 1969Jakobi, H. 1969. Forficatocaris noodti n. gen., n. sp. (Copepoda Harpacticoida) aus brasilianischem Limnopsammal. Crustaceana, 17: 231-238., MurunducarisReid, 1994Reid, J.W. 1994. Murunducaris juneae, new genus, new species (Copepoda, Harpacticoida, Parastenocarididae) from a wet campo in central Brazil. Journal of Crustacean Biology, 14: 771-781., Brasilibathynellocaris, Iticocaris, Eirinicaris, and maybe Remaneicaris (see discussion about Parastenocaris ahaggaricaBozic, 1978Bozic, B. 1978. Une nouvelle Parastenocaris du Hoggar (Crustacea, Copepoda, Harpacticoida). Bulletin du Muséum National d’Histoire Naturelle, Paris, 514: 239-244. in Corgosinho et al., 2017aCorgosinho, P.H.C.; Mercado-Salas, N.; Martínez Arbizu, P.; Santos Silva, E.N. and Kihara, T.C. 2017a. Revision of the Remaneicaris argentina-group (Copepoda, Harpacticoida, Parastenocarididae): supplementary description of species, and description of the first semi-terrestrial Remaneicaris from the tropical forest of Southeast Mexico. Zootaxa, 4238: 499-530.) are endemic to the Neotropical region. Siolicaris appears as the single genus with a hypothetical Gondwanian distribution. Most of the Neotropical genera are in Fontinalicaridinae, whereas Eirinicaris and Remaneicaris are the only representatives of Parastenocaridinae present in the region. However, specimens belonging to the monophyletic genus Parastenocaris, as redefined by Karanovic and Lee (2012Karanovic, T.; Cho, J-L. and Lee, W. 2012. Redefinition of the parastenocaridid genus Proserpinicaris (Copepoda: Harpacticoida), with description of three new species from Korea. Journal of Natural History, 46: 25-26.), were found in samples from Mongaguá (state of São Paulo) and Morretes (state of Paraná), southern Brazil (unpublished data).
In this work we describe a new genus and species of Fontinalicaridinae closely related to Murunducaris. Parastenocaris santaremensisNoodt, 1963Noodt, W. 1963. Subterrane Crustaceen der zentralen Neotropis. Zur Frage mariner Relikte im Bereich des Rio Paraguay - Paraná - Amazonas Systems. Zoologischer Anzeiger, 171: 114-147. is redescribed and designated as the type species of the new genus. The phylogenetic position of Murunducaris and Santaremicaris gen. nov. within Fontinalicaridinae is still unsolved, pending a further robust phylogenetic analysis. However, in this work we discuss the phylogenetic relationships of Murunducaris and Santaremicaris gen. nov. with other genera and species within and outside the Neotropical region.
MATERIAL AND METHODS
The type material of Santaremicaris santaremensis (Noodt, 1963Noodt, W. 1963. Subterrane Crustaceen der zentralen Neotropis. Zur Frage mariner Relikte im Bereich des Rio Paraguay - Paraná - Amazonas Systems. Zoologischer Anzeiger, 171: 114-147.) comb. nov. and S. amazonensis sp. nov. were obtained from the Noodt Collection deposited at the DZMB (Deutsches Zentrum für Marine Biodiversitätsforschung)/Senckenberg am Meer, Wilhelmshaven, Germany. The material of both species was collected in the hyporheic zone of the Amazonas River at Santarém, state of Pará, Brazil (Fig. 1). Several specimens (dissected or undissected), not always belonging to the same species, were mounted on single slides. It was often not possible to identify which limbs were previously illustrated by Noodt. As a result of the poor condition of some original slides, the redescription of S. santaremensis comb. nov. had to be based on several specimens from different slides. The description of S. amazonensis sp. nov. was based on a dissected male and an intact male mounted on a single slide along with a dissected male of S. santaremensis comb. nov, a dissected female and an intact female of S. santaremensis comb. nov, and a juvenile. The position of the voucher specimens of S. amazonensis sp. nov. are marked on the slide. Noodt’s collection has not received museum accession numbers yet, but it is organized in “slide storage maps”, prepared by Noodt, and available at the DZMB Senckenberg am Meer Wilhelmshaven (Germany). The identification of each slide was based on a reference to the “slide storage maps” (M), their sequential number and the number of the slides (preparations) within each map. Hence, for example, M2-6 is the sixth slide stored on the second slide storage map of Noodt’s collection.
Map indicating the type locality (Santarém, Pará, Brazil) of Santaremicaris santaremensis comb. Nov., and Santaremicaris amazonensis sp. nov. Arrows mark the position of Santarém within Brazil and Amazon.
Drawings were prepared on a Leica DMR microscope, fitted with Nomarski interference contrast optics and a drawing tube, at 400× and 1000× magnification. The lengths of the specimens were measured from the tip of the rostrum to the posterior rim of furca. Habitus and sensilla distribution are illustrated based on specimens picked from unsorted samples collected by Noodt at the same type locality, and stored at the DZMB.
The diagnosis represents the reconstructed ground pattern of Santaremicaris gen. nov. The term ground pattern is used in the sense of “grundmuster” (Ax, 1984Ax, P. 1984. Das Phylogenetische System. Stuttgart, Gustav Fischer Verlag, 349p., p. 156) and refers to all plesiomorphies and autapomorphies present in the stem species of the genus in question. A table with the characters differing between the species of Santaremicaris gen. nov. is included for reference (Tab. 1).
Comparative list of characters for the males of Santaremicaris santaremensis comb. nov. and Santaremicaris amazonensis sp. nov.
The terms ‘furca’ and ‘telson’ are used according to Schminke (1976Schminke, H.K. 1976. The ubiquitous telson and the deceptive furca. Crustaceana, 30: 292-300.). For homology of the furcal setae I-III we have compared our specimens with other Neotropical species belonging to Brasilibathynellocaris, Murunducaris, and Siolicaris, which all have a reduced or diminutive seta II between the larger seta I and III (see Corgosinho et al., 2008Corgosinho, P.H.C.; Martínez Arbizu, P. and Reid, J.W. 2008. Revision of the genus Murunducaris (Copepoda: Harpacticoida: Parastenocarididae), with descriptions of two new species from South America. Journal of Crustacean Biology, 28: 700-720.; 2010aCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2010a. Revision of Brasilibathynellocaris Jakobi, 1972 (Copepoda: Harpacticoida: Parastenocarididae) with redefinition of the genus. Zoological Journal of the Linnean Society, 159: 527-566., 2012bCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2010b. Three new interstitial species of Remaneicaris Jakobi, 1972 (Copepoda: Harpacticoida: Parastenocarididae) from southern and southeastern Brazil. Invertebrate Zoology, 7: 1-28.). Terminology and homology of maxillary structures follow Ferrari and Ivanenko (2008Ferrari, F.D. and Ivanenko, V.N. 2008. The identity of protopodal segments and the ramus of maxilla2 of copepods (Copepoda). Crustaceana, 81: 823-835.). Therefore, by the application of serial homology, the nomenclature of Huys and Boxshall (1991Huys, R. and Boxshall, G.A. 1991. Copepod evolution. London, The Ray Society, 468p. ) for Mx2 (Fig. 1.5.5, p. 26) is modified as follows: praecoxa of Mx2 is hereafter recognized as syncoxa (praecoxa and coxa), coxa is considered as the basis, and the basis is recognized as the first endopodal segment with claw. Other morphological terms are used according to Huys and Boxshall (1991Huys, R. and Boxshall, G.A. 1991. Copepod evolution. London, The Ray Society, 468p. ).
Abbreviations used: A1 = antennule, A2 = antenna, Ae = aesthetasc, Ap = apomorphy, Aop = anal operculum, Cph = cephalothorax, enp = endopod, exp = exopod, enp1−3 = endopodal segments 1−3, exp1−3 = exopodal segments 1−3, Fig. = figure, Figs.= Figures, Fu = furca, Gds = genital-double somite, Gf = genital field, Ms = modified seta, Md = mandible, Mf = modified spine, Mx1 = maxillule, Mx2 = maxilla, Mxp = maxilliped, P1−P5 = legs 1 to 5, Pl = plesiomorphy, Sy = synapomorphy, Ur(s) = urosomite(s), Ur1−5 = urosomites 1−5.
SYSTEMATICS
Harpacticoida Sars, 1903Sars, G.O. 1903. An account of the Crustacea of Norway, 5 Copepoda Harpacticoida. Parts I and II, 1-28.
Parastenocarididae Chappuis, 1940Chappuis, P.A. 1940. Die Harpacticoiden des Grundwassers des unteren Maintales. Archiv für Hidrobiologie, 36: 286-305.
Fontinalicaridinae Schminke, 2010Schminke, H.K. 2010. High-level phylogenetic relationships within Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 83: 343-367.
Santaremicaris gen. nov.
ZooBank: urn:lsid:zoobank.org:act:27524A69-87E4-4634-A626-62D514F995A1
Diagnosis. Small-sized Fontinalicaridinae (255−325 μm), with cylindrical habitus; body cuticle poorly chitinised; dorsal integumental window on Cph, Urs 3−5 in male and Urs 2−4 in female; boundary between prosome and urosome inconspicuous; Gf a wide concave plate, two times wider than narrow, with medial process and occupying anterior ventral half of female Gds. Last Urs of males and females with undulate distal frill. Fu cylindrical, divergent, about three times as long as wide, armed with six setae (Setae I, III, IV, V, VI, and VII), with distinct gap between setae I−III and VII, setae I and III inserted in proximal third of furca, seta VII in distal third; seta I longer than seta III, seta II absent, seta IV distally inserted on outer margin, longer than setae I, III, VI and VII, seta V the longest, about 4 times longer than Fu. Antennule eight-segmented in male and seven-segmented in female. A1 of the coiled type, segments V−VIII of males forming a functional unit for clasping the female. Coxal gnathobasis of md with distal row of teeth and seta; md palp one-segmented, with two distal setae. Mx1 praecoxal arthrite with five elements (surface seta thick and blunt, three claw-like pinnate spines, and slender seta); coxal endite with one seta; basis with three setae. Mx allobasis with two endites; proximal endite with one seta; distal endite with two smooth setae and serrate spine; proximal endopodal segment drawn out into claw; distal endopodal segment with two setae. Mxp subchelate; syncoxa about 1/3 length of basis; enp drawn into spinulose claw. Basis of P1 sexually dimorphic in male, with modified robust spinule or row of strong spinules on inner margin; with outer seta. Male P2 enp with long distal seta, the insertion area of it covering almost the whole width of the segment, without distal ornamentation and with strong ornamentation on outer margin, along the first 2/3 of the segment. Male P3 with rectangular, smooth or ornamented coxa; basis rectangular or trapezoidal, approximately twice as long as wide, with long row of spinules on inner margin; with hook-like spinule on proximal inner margin or without it; enp absent; exp one-segmented, rectangular with regular margins or with irregular margins and tapering distally, longer than wide, slightly curved inwards, weakly ornamented on outer margin or smooth, with or without tube pore on inner margin; apophysis without distal seta or spine, spatuliform and shorter than thumb or slightly longer than thumb, the apophysis ending in a long tip with a rounded hyaline cushion at tip; thumb with or without hyaline inner margin. Female P3 enp unarmed, shorter than exp1 and acuminate distally. Male P4 basis unornamented, with outer seta and outer pore; inner margin of male P4 exp1 with distal and proximal row of spinules; enp short, triangular, not reaching middle of exp1, smooth or ornamented on outer margin, with strongly chitinized regions. Female P4 enp one-segmented, slightly longer than exp1 and bearing one distal and bipinnate spine fused to segment, ornamented with spinules on distal half, near the fusion of the spine and the enp. Male P5 a well-developed triangular plate reaching middle of next somite or slightly beyond; with irregular margins; inner margin with row of small spinules; distal portion ending in strong spiniform process; armature consisting of very long outer basal seta and two additional setae; intercoxal sclerite enlarged, triangular or a trapezoid plate, longer than wide, with distomedial cuticular process. Female P5 with small distal spiniform process, shorter than in male, long outer seta and three well-developed lateral setae; with a small triangular intercoxal sclerite. P6 an unarmed concave plate with median hump covering Gf.
Type species. Parastenocaris santaremensisNoodt, 1963Noodt, W. 1963. Subterrane Crustaceen der zentralen Neotropis. Zur Frage mariner Relikte im Bereich des Rio Paraguay - Paraná - Amazonas Systems. Zoologischer Anzeiger, 171: 114-147..
Composing species. Santaremicaris santaremensis (Noodt, 1963Noodt, W. 1963. Subterrane Crustaceen der zentralen Neotropis. Zur Frage mariner Relikte im Bereich des Rio Paraguay - Paraná - Amazonas Systems. Zoologischer Anzeiger, 171: 114-147.) comb. nov.; S. amazonensis sp. nov.
Etymology. The generic name refers to Santarém (Pará, Brazil), the municipality in which the new species were collected, combining this name with the ancient Greek substantive for shrimp, καρίς (caris).
Autapomorphies for the genus. Male second urosomite without integumental window. Last Ur of males and females with undulate distal frill. P2 enp without distal ornamentation, with long distal seta, the insertion area of it covering almost the whole width of the segment, and strong outer ornamentation along the first 2/3 of the segment. Basis of male P3 with long row of spinules on the inner margin. Absence of male P3 enp. Short, triangular and strongly chitinized male P4 enp. Long and slim male P5 with irregular margins, reaching the middle of the next urosomite or beyond, distal portion ending in a strong spiniform process, ornamented with spinules on the inner margin, armed with only three setigerous elements on the outer margin.
Santaremicaris santaremensis ( Noodt, 1963Noodt, W. 1963. Subterrane Crustaceen der zentralen Neotropis. Zur Frage mariner Relikte im Bereich des Rio Paraguay - Paraná - Amazonas Systems. Zoologischer Anzeiger, 171: 114-147. ) comb. nov.
Santaremicaris santaremensis comb. nov., male. Antennule (A), antenna (B), and last urosomite, telson, and furca (C). Roman numerals identify the antennule segments and furcal setae. Vertical scale bar (A, B), horizontal scale bar (C).
Santaremicaris santaremensis comb. nov., male. P1 (A), P2 (B), P3 (C), P4 (D), and P5 (E).
Santaremicaris santaremensis comb. nov., female. Dorsal (A), lateral (B) habitus, and left furca in dorsal view (C). Roman numerals identify furcal setae. Vertical scale bar (A, B), horizontal scale bar (C).
Santaremicaris santaremensis comb. nov., female. Antennule (A and B), mandible (C), maxillula (D), maxilla (E), and maxilliped (F). Roman numerals identify the antennule segments.
Santaremicaris santaremensis comb. nov., female. P1 (A), protopodite, exp1, and enp of P2 (B), P3 (C), protopodite, exp1, and enp of P4 (D), P5 (E).
Santaremicaris santaremensis comb. nov., female. First urosomite with P5 and genital double-somite with genital field.
Type material. M5-6. Slide containing two dissected males and some females.
Other examined material. M2-6; M2-8; M5-5 and undissected male and female picked from an unsorted sample collected by Noodt, at the same type location of S. santaremensis comb. nov. and stored at the DZMB. See material and methods for explanation of the codes.
Type locality. Hyporheic zone of the Amazonas River at Santarém (Pará, Brazil) (Noodt, 1963Noodt, W. 1963. Subterrane Crustaceen der zentralen Neotropis. Zur Frage mariner Relikte im Bereich des Rio Paraguay - Paraná - Amazonas Systems. Zoologischer Anzeiger, 171: 114-147.) (Fig. 1). Approximate coordinates: 02°25’S 54°42’W.
Other occurrences. Upper reaches of Negro River, lower reaches of the Tapajós and Tocantins Rivers.
Description of male (Figs. 2−4). Total length 285 µm (much shorter than measures given by Noodt, 1963Noodt, W. 1963. Subterrane Crustaceen der zentralen Neotropis. Zur Frage mariner Relikte im Bereich des Rio Paraguay - Paraná - Amazonas Systems. Zoologischer Anzeiger, 171: 114-147., p. 144: 370−380µm), measured from rostrum to end of furca. Rostrum not fused to cephalothorax, with wide base and two sensilla on tip (Fig. 2A, B). Cephalothorax and Urs 3−5 with dorsal integumental window (Fig. 2A, B), absent on Ur2. Pattern of sensilla as depicted (Fig. 2A, B); last Ur with distal frill undulate and thicker than those of previous Urs (Figs. 2A, B, 3C). Telson smooth (Fig. 3C), with proximal pore on lateral margins, with pair of sensilla adjacent to anal operculum, and without additional ornamentation; Aop slightly concave medially (Fig. 2A); Fu (Figs. 2A, B, 3C) cylindrical, approximately 2.5 times as long as wide, slightly enlarged distally in lateral view, slightly irregular on outer margin and convex on inner margin in dorsal view; presence of distal outer pore, and two lateral setae on proximal third; seta I longer than seta III, almost as long as setae VI and VII (Fig. 3C); seta II absent; seta IV distally inserted on outer margin, longer than setae I, III, VI and VII; seta V the longest, about four times longer than Fu; dorsal seta (VII) at distal third; all setae smooth except for unipinnate seta IV.
A1 (Fig. 3A): eight-segmented; of the coiled type with segments V to VIII forming functional unit for clasping female; armature as follows: I(0)/II(6)/III(4)/IV(1+1Ms)/V(4+(1+Ae))/VI(2+Ms)/VII(1+Ms)/VIII(7+ acrothec of 2+Ae).
A2 (Fig. 3B): allobasis without abexopodal armature, with small outer spinule proximally; one-segmented exp with long unipinnate seta; free endopodal segment with two proximal rows of spinules on inner margin, and transverse row of spinules subterminally; presence of two inner marginal spines and five terminal setae/spines, two of them geniculated.
Labrum triangular in lateral view.
Md: Coxal gnathobasis with distal row of teeth and seta; palp one-segmented, with two distal setae.
Mx1: Praecoxal arthrite with five elements (surface seta thick and blunt, three claw-like pinnate spines, and slender seta); coxal endite with one seta; basis with three setae.
Mx: Allobasis with two endites; proximal endite with one seta; distal endite with two smooth setae and serrate spine; proximal endopodal segment drawn out into claw; distal endopodal segment with two setae.
Mxp: subchelate; syncoxa about 1/3 length of basis; enp drawn into spinulose claw.
P1 (Fig. 4A): coxa smooth. Anterior surface of basis with row of spinules on inner margin, distalmost enlarged; rows of spinules near insertion of enp and exp on anterior surface; and outer seta, outer pore and two outer spinules. Enp two-segmented: enp1 nearly as long as exp1 and exp2 combined, with two inner rows, two outer rows and one distal row of spinules (Fig. 4A); enp2 with outer spine and long geniculate seta, with posterior hyaline frill. Exp three-segmented: exp1 with outer spine, rows of spinules on outer margin, proximal and distal to outer spine; exp2 unarmed, with row of outer spinules along outer margin; exp3 with outer row of spinules proximally, two outer spines and two geniculate apical setae.
P2 (Fig. 4B): intercoxal sclerite a small unornamented trapezoidal plate, shorter than coxa, concave proximally and distally; with proximal lateral projections. Coxa with anterior row of spinules. Basis without outer seta, with row of spinules on outer margin and outer pore on anterior surface. Enp one-segmented, 0.4 times as long as exp1, with distal seta as long as exp1, the insertion area of it covering almost the whole width of the segment; with strong spinules on outer margin, not reaching the distal 1/3, distal margin unornamented. Exp three-segmented: exp1 with outer row of spinules, hyaline frill distally on inner corner, and outer spine reaching beyond distal edge of exp2; exp2 with distal row of strong spinules only; exp3 with outer distal row of spinules, hyaline frill at inner distal corner, unipinnate distal outer spine, unipinnate distal medial seta and distal inner seta (distal inner seta distally broken).
P3 (Fig. 4C): intercoxal sclerite a small unornamented trapezoidal plate as long as coxa, straight proximally, concave distally, with proximal lateral projections. Coxa rectangular, with diagonal row of spinules on anterior face. Basis long, trapezoidal, bearing long outer seta and outer pore, hook-like spinule on proximal inner corner and row of spinules. Enp lacking. Exp one-segmented, rectangular, long, slightly curved inwards, smooth on outer margin, with long and spatuliform apophysis; thumb longer than apophysis, with medial hump and ending in long tip.
P4 (Fig. 4D): intercoxal sclerite a small unornamented trapezoidal plate, shorter than coxa, straight proximally, concave distally, with proximal lateral projections. Coxa smooth. Basis not ornamented, with outer pore on anterior surface; outer seta long, reaching beyond distal margin of exp1. Enp triangular, with strongly chitinized areas, short, approximately 0.7 times as long as exp1, smooth and acuminate. Exp three-segmented: exp1 with outer row of spinules in proximal third; rows of spinules proximally and distally to outer spine reaching distal edge of exp2; inner margin with row of spinules from median to distal margin replacing hyaline frill, with additional row of spinules in proximal third; exp2 with distal row of strong spinules; exp3 with distal row of spinules on outer margin, inner hyaline frill at distal corner, unipinnate outer spine, and distal unipinnate seta nearly twice as long as outer seta.
P5 (Fig. 4E): intercoxal sclerite a large triangular plate reaching the proximal 1/3 of the limb, with distomedial claviform cuticular process. P5 consisting of well-developed triangular plate reaching middle of subsequent Ur; proximal 1/3 with irregular inner margin and outer hump, medially with concave inner margin and convex outer margin; distally with convex inner margin and concave outer margin; inner margin bearing row of small spinules; distal portion spiniform; armature consisting of very long outer basal seta and 2 additional setae;
P6 represented by unarmed cuticular flap.
Description of female (Figs. 5−8). Length 351μm (slightly shorter than measurements given by Noodt, 1963Noodt, W. 1963. Subterrane Crustaceen der zentralen Neotropis. Zur Frage mariner Relikte im Bereich des Rio Paraguay - Paraná - Amazonas Systems. Zoologischer Anzeiger, 171: 114-147., p. 144: 350−380μm), measured from tip of rostrum to end of furca. Sexual dimorphism expressed in A1, P1, P2, P3, P4, P5 and Gds. Cph and Urs2 (Gds), 3, and 4 with dorsal integumental windows. Urs2−4 with undulate distal frill on dorsal view, Ur4 with undulate distal frill on lateral view. Telson as in male, except for the more concave Aop (Fig. 5A). Fu (Fig. 5A−C) cylindrical, approximately 2.5 times longer than wide, slightly enlarged distally in lateral view, convex on inner margin in dorsal view, concave in the outer margin, with distal outer pore, and two lateral setae at end of proximal third. Fu armature as follows: seta I and III subequal, both about twice shorter than setae IV and VII; seta II absent; seta IV almost as long as setae VI and VII, subterminally inserted on outer margin; seta V the longest, 3.4 times longer than Fu; dorsal seta (VII) at distal third; all setae smooth, except for unipinnate setae IV and VII.
A1 seven-segmented (Fig. 6A, B); armature as follows: I(0)/II(4)/III(4)/IV(1+ (1+ae))/V(0)/VI(0)/VII(7+ (2+ae)).
A2 as in male.
Md (Fig. 6C), Mx1 (Fig. 6D), Mx (Fig. 6E), Mxp (Fig. 6F) as described for male.
P1 (Fig. 7A): intercoxal sclerite a small unornamented trapezoidal plate, shorter than coxa, concave proximally and distally. Coxa as in male; basis with row of spinules on inner margin not as developed as in male, without row of spinules near insertion of enp and exp, and with five outer spinules; exp1−3 as in male; enp1 lacking proximal row of spinules on inner margin, enp2 as in male.
P2 (Fig. 7B): intercoxal sclerite a small unornamented trapezoidal plate, shorter than coxa, slightly concave proximally and distally, with proximal lateral projection. Coxa smooth; basis with row of spinules near insertion of enp, outer pore and outer row of spinules; outer margin of exp1−3 as in male; enp thinner than in male, without outer ornamentation along the proximal 2/3, with subdistal spinules, two on inner margin, one on outer margin, with thin and long distal seta, as long as exp1, the insertion area not as wide as in the male.
P3 (Fig. 7C): intercoxal sclerite a small unornamented trapezoidal plate, shorter than coxa, proximally straight and irregular distally. Coxa smooth; basis with outer pore, row of spinules distal to long outer seta, and row of spinules near insertion of enp; exp two-segmented, exp1 enlarged, with strong spinules in proximal third, proximally and distally to outer spine, distal row of strong spinules and row of small spinules on inner margin; exp2 with subdistal row of spinules on outer margin, with inner hyaline frill, outer bipinnate spine and distal unipinnate seta; enp one-segmented, 0.5 times as long as exp1, unarmed, acuminate distally .
P4 (Fig. 7D): intercoxal sclerite a small unornamented trapezoidal plate, shorter than coxa, almost straight proximally, concave distally, with proximal lateral projection. Coxa smooth; basis with row of spinules distally to outer seta, and row of spinules near enp insertion; exp1 without ornamentation on inner margin and with a distal inner hyaline frill, outer margin with row of spinules in proximal third, proximally and distally to outer spine; exp2−3 as in male; P4 enp one-segmented, slightly longer than exp1 and bearing one distal and bipinnate spine fused to segment; with two spinules on inner margin, medial and anterior spinules inserted at base of spine.
P5 (Fig. 7E−8): well developed, a simple trapezoidal plate; distal margin ending on spiniform process, without inner ornamentation, reaching distal margin of the Gds; armature composed of proximal, long and articulated seta, two medial setae, and distal seta. Intercoxal sclerite a small triangular plate.
P6 (Fig. 8): unarmed concave plate with median hump covering Gf.
Santaremicaris amazonensis sp. nov.
ZooBank: urn:lsid:zoobank.org:act:976B796E-DE41-494F-BC5D-21BCD727878B
Santaremicaris amazonensis sp. nov., male. Dorsal (A), lateral (B) habitus, and right furca in dorsal view (C). Roman numerals identify furcal setae. Vertical scale bar (A, B), horizontal scale bar (C).
Type material. Only one intact male and one dissected male of S. amazonensis sp. nov. present in the slide M2-6 are designated as the syntypes. The dissected male of S. santaremensis comb. nov., the intact and the dissected female of S. santaremensis comb. nov., and the juvenile mounted in the same slide are not part of the type series.
Other examined material. Undissected male picked from an unsorted sample collected by Noodt, at the same type location of P. santaremensis comb. nov. and stored at the DZMB. See material and methods for explanation of the codes.
Type locality. Hyporheic zone of the Amazonas River at Santarém (Pará, Brazil) (Fig. 1). Approximate coordinates: 02°25’S 54°42’W.
Etymology. The specific name refers to the Amazonian biome in which the new species was found.
Description of male (Figs. 9, 10). Length 351µm, measured from rostrum to end of furca. Rostrum not fused to cephalothorax, with wide base and two sensilla on tip (Fig. 9A, B). Cephalothorax and Urs3−5 with dorsal integumental window (Fig. 9A, B), last Ur with undulate distal frill. Patterns of sensilla as depicted (Fig. 9A, B). Telson with pair of sensilla adjacent to anal operculum, without additional ornamentation and with proximal lateral pore. Fu cylindrical (Fig. 9A−C), about 2.5 times as long as wide, slightly enlarged distally in lateral view, slightly irregular on outer margin and slightly convex on inner margin in dorsal view, with distal outer pore, and two lateral setae in proximal third. Armature of Fu as follows (Fig. 9A−C): seta I shorter than seta III, seta II absent, seta III shorter than seta IV; seta IV inserted distally on outer margin, almost as long as seta VII, dorsal seta VII inserted at distal third, longer than seta VI; seta V the longest, more than 3 times longer than furca; all setae smooth.
A1 and A2 as in S. santaremensis comb. nov.
Labrum triangular in lateral view. Md, Mx1, Mx2 and Mxp as described for S. santaremensis comb. nov.
P1 (Fig. 10A): intercoxal sclerite a small unornamented trapezoidal plate, shorter than coxa, straight proximally, concave distally, with proximal lateral projection. Coxa smooth. Basis with row of strong spinules on inner margin, row of minute spinules near insertion of enp and distal to outer seta. Enp two-segmented: enp1 longer than exp1 and exp2 combined, with one inner row, two outer rows and one terminal row of spinules; enp2 with outer spine, long geniculate seta, and posterior hyaline frill. Exp three-segmented: exp1 bearing outer spine, proximal and distal rows of spinules on outer margin; exp2 unarmed, with row of outer spinules; exp3 with outer row of spinules proximally, two outer spines and two geniculate apical setae.
P2 (Fig. 10B): as in S. santaremensis comb. nov., except for presence of spinules on anterior surface of basis, near enp insertion, and stronger spinules on outer margin of enp.
P3 (Fig. 10C): Coxa quadratic. Basis rectangular, with long outer seta, outer pore, and V-shaped row of spinules on inner margin. Enp lacking. Exp unisegmented, irregular in shape, long, slightly curved inwards, with cuticular humps on medial and proximal inner margin, inner tube pore near articulation with basis, and with two proximal outer spinules; apophysis long, sigmoid, narrowing in long tip with distal hyaline cushion; thumb shorter than apophysis, with long tip and hyaline inner margin.
P4 (Fig. 10D): similar to S. santaremensis comb. nov., except for presence of row of spinules on outer margin of enp; stronger proximal row of spinules on exp1 inner margin, and weaker distal row of spinules on exp1 inner margin.
P5 (Fig. 10E): intercoxal sclerite a large trapezoidal plate reaching the proximal 1/3 of the limb, with distomedial linguiform cuticular process. P5 consisting of well-developed triangular plate reaching middle of subsequent Ur; with irregular outer margin, slightly bulging proximally; inner margin almost straight, with row of small spinules; distal portion ending in strong and thin spiniform process; armature consisting of very long outer basal seta and two additional setae inserted on distalmost hump.
P6 represented by unarmed cuticular flap.
DISCUSSION
Most of the characters proposed by Schminke (2010Schminke, H.K. 2010. High-level phylogenetic relationships within Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 83: 343-367.) cannot be satisfactorily used as diagnostic for the Neotropical Fontinalicaridinae (i.e., Brasilibathynellocaris, Colombocaris, Eirinicaris, Forficatocaris, Iticocaris, Murunducaris, Noodtcaris, Potamocaris, Remaneicaris, Siolicaris).
According to Schminke (2010Schminke, H.K. 2010. High-level phylogenetic relationships within Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 83: 343-367.), in the Fontinalicaridinae, a) the terminal seta of the male P3 is separate from the apophysis; b) the enp of female P3 has no terminal seta and is a relatively short element with a rounded apex, carrying spinules terminally and subterminally; c) the basis of male P4 has an inner row of spinules and is ornate between the exp and enp insertions; d) the P5 of both sexes are much larger and triangular, and extend back well beyond the own somite; and e) in the furca, the group of three lateral setae and the dorsal seta are at a distance from one another, the dorsal seta being located distally, the three lateral setae proximally.
However, within the Neotropical Fontinalicaridinae the unfused spine on male P3 apophysis is present in Noodtcaris kubitzkii (Noodt, 1972Noodt, W. 1972. Brasilianische Grundwasser-Crustacea, 1. Studien an den Gattungen Parastenocaris Kessler und Forficatocaris Jakobi aus der Serra do Mar von São Paulo (Copepoda, Harpacticoida). Crustaceana, 23: 76-99.) and Noodtcaris tapajoensis (Noodt, 1963Noodt, W. 1963. Subterrane Crustaceen der zentralen Neotropis. Zur Frage mariner Relikte im Bereich des Rio Paraguay - Paraná - Amazonas Systems. Zoologischer Anzeiger, 171: 114-147.), but it is modified in these species as a hyaline spine in N. kubitzkii or it is a blunt spine in N. tapajoensis. In the remaining Noodtcaris species, the spine is fused to the apophysis. The referred spine is not fused to the apophysis in Potamocaris and Forficatocaris, but its loss as an articulated element or fusion to the apophysis occurs within Potamocaris. The modification of the spine into a hyaline spine occurs in some species of Potamocaris and Forficatocaris. Only Murunducaris has the typical unfused spine, although strongly modified in Murunducaris noodtiCorgosinho, Martínez Arbizu and Reid, 2008Corgosinho, P.H.C.; Martínez Arbizu, P. and Reid, J.W. 2008. Revision of the genus Murunducaris (Copepoda: Harpacticoida: Parastenocarididae), with descriptions of two new species from South America. Journal of Crustacean Biology, 28: 700-720.. Siolicaris has an unarmed apophysis, with the exception of Siolicaris sandhya (Ranga Reddy, 2001Ranga Reddy, Y. 2001. Discovery of Parastenocarididae (Copepoda, Harpacticoida) in India, with the description of three new species of Parastenocaris Kessler, 1913, from the river Krishna at Vijayawada. Crustaceana, 74: 705-733.) (Corgosinho et al., 2012bCorgosinho, P.H.C.; Ranga Reddy, Y. and Martínez Arbizu, P. 2012b. Revision of the genus Siolicaris Jakobi, 1972, with redescriptions of S. sioli (Noodt, 1963) and S. jakobi (Noodt, 1963) from South America, and S. sandhya (Ranga Reddy, 2001) comb. nov. from India (Copepoda, Harpacticoida, Parastenocarididae). Zootaxa, 3493: 49-71.). The apophysis is also unarmed on Iticocaris and S. amazonensis sp. nov., but bears a distal hyaline cushion, which we consider homologous to a spine, in Brasilibathynellocaris and S. santaremensis comb. nov. The female P3 enp is absent in Potamocaris, Siolicaris and at least in the Colombian species of Noodtcaris (the female of N. tapajoensis is unknown). When the enp is present on the female P3, it is generally short, unarmed, and with blunt tip, but it is bifid on Murunducaris dactyloides (Kiefer, 1967Kiefer, F. 1967. Zwei weitere Parastenocaris-Arten (Copepoda, Harpacticoida) aus dem mittleren Amazonas-Gebiet. Amazoniana, 1: 131-134.), bears a distal seta on Murunducaris loyolaiCorgosinho, Martínez Arbizu and Reid, 2008Corgosinho, P.H.C.; Martínez Arbizu, P. and Reid, J.W. 2008. Revision of the genus Murunducaris (Copepoda: Harpacticoida: Parastenocarididae), with descriptions of two new species from South America. Journal of Crustacean Biology, 28: 700-720., and has an acuminate tip in S. santaremensis comb. nov. The basal accessory spinule or a row of spinules between enp and exp of male P4 is generally present in Potamocaris and Forficatocaris. This character is absent in Murunducaris juneaeReid, 1994Reid, J.W. 1994. Murunducaris juneae, new genus, new species (Copepoda, Harpacticoida, Parastenocarididae) from a wet campo in central Brazil. Journal of Crustacean Biology, 14: 771-781., Colombocaris isabellaeGaviria, Defaye and Corgosinho, 2017Gaviria, S.; Defaye, D. and Corgosinho, P.H.C. 2017. Two new parastenocaridid genera (Copepoda, Harpacticoida) from South America, including a revision of Noodt’s columbiensis-group and the redescription of Parastenocaris tapajoensis Noodt, 1963. Crustaceana, 90: 535-570. , Noodtcaris columbiensis (Noodt, 1972Noodt, W. 1972. Brasilianische Grundwasser-Crustacea, 1. Studien an den Gattungen Parastenocaris Kessler und Forficatocaris Jakobi aus der Serra do Mar von São Paulo (Copepoda, Harpacticoida). Crustaceana, 23: 76-99.), Brasilibathynellcaris, Siolicaris partim and Iticocaris. The P5 morphology shows a plethora of transformations in shape, size, and armature within the Neotropical Fontinalicaridinae. It is dimorphic in shape and armature in Potamocaris, Forficatocaris, Murunducaris, and Santaremicaris gen. nov. (maybe also in Colombocaris, but the female is unknown), with extreme and non-homologous transformations occurring in Murunducaris and Forficatocaris. The P5 is rather short in Siolicaris and Brasilibathynellocaris. The typical fontinalicaridid P5 as described by Schminke (2010Schminke, H.K. 2010. High-level phylogenetic relationships within Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 83: 343-367.) is present in Iticocaris and Noodtcaris only. The most consistent diagnostic character is the insertion of the lateral setae of the furca (setae I, II, III) anteriorly to the dorsal one (seta VII). This character differs only in Potamocaris, where the lateral and dorsal setae are distally inserted, at the same transverse plane.
Therefore, it is evident that most of the diagnostic characters are very plastic within Neotropical fontinalicaridids, and the inclusion of the new genus within this subfamily must be on the grounds of derived characters shared with Murunducaris plus the insertion of the lateral furcal setae anteriorly to the dorsal one, and the presence of a short, not spiniform, and unarmed female P3 enp.
A robust phylogenetic hypothesis on the position of the monophylum Murunducaris/Santaremicaris gen. nov. within Fontinalicaridinae is not possible without a broader morphological and molecular study with sufficient taxon sampling. However, some evidence indicates a closer proximity between Potamocaris, Forficatocaris, Murunducaris, Santaremicaris gen. nov., Parastenocaris fossorisFryer, 1956Fryer, G. 1956. New species of cyclopoid and harpacticoid copepods from sandy beaches of Lake Nyasa. Annals and Magazine of Natural History, 12: 225-249., Parastenocaris crassicaudisChappuis, 1955Chappuis, P.A. 1955. Harpacticoïdes psammiques du lac Tanganika. Revue de Zoologie et de Botanique Africaines, 51: 68-80., Parastenocaris nigerianusChappuis, 1959Chappuis, P.A. 1959. Deux Parastenocaris des sables du Niger. Bulletin de l’Institut Fondamental d’Afrique Noire, Série A, Sciences Naturelles, 21:1211-1215., Parastenocaris jeannineiDumont, 1981Dumont, H.J. 1981. Cladocera and free-living Copepoda from the Fouta Djalon and adjacent mountain areas in West Africa. Hydrobiologia, 85:97-116., Parastenocaris kimiDumont, 1981Dumont, H.J. 1981. Cladocera and free-living Copepoda from the Fouta Djalon and adjacent mountain areas in West Africa. Hydrobiologia, 85:97-116., Parastenocaris aberransApostolov, 2004Apostolov, A. 2004. Copépodes harpacticoïdes stygobies de France. 6. Description d’un nouveau représentant du genre Parastenocaris Kessler, 1913. Crustaceana, 77: 197-203., and maybe Proserpinicaris phyllura (Kiefer, 1938Kiefer, F. 1938. Neue harpacticoide Ruderfusskrebse (Crust. Cop.) aus dem Grundwasser von Aschaffenburg (Main). Zoologischer Anzeiger, 123(5-6): 142-147.) (incertae sedis). For details, see discussion below.
Characters shared between Santaremicaris gen. nov. and Murunducaris.
The new genus shares with Murunducaris (1) the presence of strong ornamentation on the inner margin of the basis of P1 in males (Sy), (2) the presence of a large male P5 intercoxal sclerite (Sy) with medial cuticular process (Sy), and (3) a proximal row of spinules on the inner margin of the male P4 exp1 (Sy). The characters are described for the two genera in details below:
(1) Strong ornamentation on the inner margin of the basis of P1 in males. The presence of an inner spine on the basis of P1 is a widespread character of Harpacticoida (Huys and Boxshall, 1991Huys, R. and Boxshall, G.A. 1991. Copepod evolution. London, The Ray Society, 468p. ; Willen, 2000Willen, E. 2000. Phylogeny of the Thalestridimorpha Lang, 1944 (Crustacea, Copepoda). Cuvillier Verlag, Göttingen . 233p.; Seifried, 2003Seifried, S. 2003. Phylogeny of Harpacticoida (Copepoda): revision of ‘Maxillipedasphalea’ and Exanechentera. Cuvillier Verlag, Göttingen. 259p.). Within Harpacticoida a strongly dimorphic spine is an apomorphy for at least the Ameiridae Boeck, 1865Boeck, A. 1865. Oversigt over de ved Norges Kyster jagttagne Copepoder henhörende til Calanidernes, Cyclopidernes og Harpactidernes Familier. Forh Vidensk Selsk Krist, 1864: 226-282. and the Diosaccinae Sars, 1906Sars, G.O. 1906. Copepoda Harpacticoida. Parts XI & XII. Thalestridae (continued), Diosaccidae (part). An Account of the Crustacea of Norway, with short descriptions and figures of all the species. Bergen Museum, 5: 133-156.. One characteristic of armature elements such as setae and spines is their hollow core, in which a single nerve enters (Fahrenbach, 1962Fahrenbach, W. H. 1962. The biology of a harpacticoid copepod. La Cellule, 62: 303-376.). Setules, spinules and cilia are solid projections of the cuticle, lacking cellular components (Fahrenbach, 1962Fahrenbach, W. H. 1962. The biology of a harpacticoid copepod. La Cellule, 62: 303-376.), and they do not cross the cuticle; this seems to be the condition present in Murunducaris and S. santaremensis comb. nov. In all Murunducaris species there is a strong and long spinule, sometimes followed by smaller accompanying spinules on the inner margin of the basis of the male P1 (Corgosinho et al., 2008Corgosinho, P.H.C.; Martínez Arbizu, P. and Reid, J.W. 2008. Revision of the genus Murunducaris (Copepoda: Harpacticoida: Parastenocarididae), with descriptions of two new species from South America. Journal of Crustacean Biology, 28: 700-720.). The same character is observed in S. santaremensis comb. nov. However, in S. amazonensis sp. nov., there is a row of strong and subequal spinules on the inner margin of the male P1. In a series of transformation, we consider that a row of long spinules as present in S. amazonensis sp. nov. precedes the condition present in S. santaremensis comb. nov. and Murunducaris; the condition in S. amazonensis sp. nov. representing the retention of a plesiomorphic condition. A weak ornamentation, with short, not enlarged spinules is observed in females of Murunducaris and S. santaremensis comb. nov. The female is unknown for S. amazonensis sp. nov. and the female P1 of M. noodti is mistakenly described as being equal to the male P1 (Corgosinho et al., 2008Corgosinho, P.H.C.; Martínez Arbizu, P. and Reid, J.W. 2008. Revision of the genus Murunducaris (Copepoda: Harpacticoida: Parastenocarididae), with descriptions of two new species from South America. Journal of Crustacean Biology, 28: 700-720.). In the female of M. noodti the basis of the P1 does not have the modified inner spinule, which is present in the males of the genus Murunducaris. We consider that this is convincing evidence supporting that the large basal spinule observed for Murunducaris and S. santaremensis comb. nov. is not homologous to the seta or spine, sometimes found in Parastenocaridinae. Within this family, when a seta or spine occurs on the inner margin of the basis of P1, it is normally present in both males and females. To our knowledge, such armature was never described for a Fontinalicaridinae species, occurring convergently only in some groups of species within the Parastenocaridinae. When a seta occurs on the inner margin of the P1 basis within Parastenocarididae, it is not dimorphic in shape, differing at most in length (e.g., Eirinicaris antonioiCorgosinho, Schizas, Previattelli, Rocha and Santos-Silva, 2017bCorgosinho, P.H.C.; Schizas, N.V.; Previattelli, D.; Rocha, C.E.F. da and Santos-Silva, E.N. 2017b. A new genus of Parastenocarididae (Copepoda, Harpacticoida) from the Tocantins River basin (Goiás, Brazil), and a phylogenetic analysis of the Parastenocaridinae. Zoosystematics and Evolution, 93: 167-187.). The absence of a modified medial spine in the male P1 basipodite is considered a synapomorphy for PsammonitocrellaHuys, 2009Huys, R. 2009. Unresolved cases of type fixation, synonymy and homonymy in harpacticoid copepod nomenclature (Crustacea: Copepoda). Zootaxa, 2183: 1-99. and Parastenocarididae (see Martínez Arbizu and Moura, 1994Martínez Arbizu, P. and Moura, G. 1994. The phylogenetic position of the Cylindropsyllinae Sars (Copepoda, Harpacticoida) and the systematic status of the Leptopontiinae Lang. Zoologische Beiträge, 35: 55-77.; Corgosinho et al., 2020Corgosinho, P.H.C.; Kihara, T.C. and Martínez Arbizu, P. 2020. A new species of Psammonitocrella Huys, 2009 (Copepoda, Harpacticoida, Ameiridae) from California (USA), with a discussion of the relationship between Psammonitocrella and Parastenocarididae. ZooKeys, 996: 19-35. ), and is a condition present in the ground pattern of the Fontinalicaridinae. Murunducaris and Santaremicaris gen. nov. are derived genera within the Fontinalicaridinae. If future studies reveal that what we consider a transformed spinule is in fact a transformed spine, it must be considered a synapomorphy for these genera, resulting from the re-expression of a plesiomorphic condition not found within the Psammonitocrella/Parastenocarididae monophylum. For additional information on this character see Corgosinho et al. (2008Corgosinho, P.H.C.; Martínez Arbizu, P. and Reid, J.W. 2008. Revision of the genus Murunducaris (Copepoda: Harpacticoida: Parastenocarididae), with descriptions of two new species from South America. Journal of Crustacean Biology, 28: 700-720.).
(2) Presence of an enlarged male P5 intercoxal sclerite with medial cuticular process. The male intercoxal sclerite of Murunducaris and Santaremicaris gen. nov. is the strongest evidence supporting the sister group relationship of these genera. This character was described for the first time by Reid (1994Reid, J.W. 1994. Murunducaris juneae, new genus, new species (Copepoda, Harpacticoida, Parastenocarididae) from a wet campo in central Brazil. Journal of Crustacean Biology, 14: 771-781.) and further confirmed by Corgosinho et al. (2008Corgosinho, P.H.C.; Martínez Arbizu, P. and Reid, J.W. 2008. Revision of the genus Murunducaris (Copepoda: Harpacticoida: Parastenocarididae), with descriptions of two new species from South America. Journal of Crustacean Biology, 28: 700-720.) for three additional species of Murunducaris and S. santaremensis comb. nov. The intercoxal sclerite is enlarged, quadratic or trapezoidal, with a cuticular process of various shapes depending on the species (e.g., hook-like, spiniform, club-shaped or linguiform) occurring between the limbs and occupying a medial or distal position in the intercoxal sclerite. This character is shared by Murunducaris and Santaremicaris gen. nov., exclusively. Further studies are necessary to verify if this character is present in Pa. fossoris and Pa. crassicaudis, two species that are considered closely related to Murunducaris and S. santaremensis comb. nov. (Corgosinho et al., 2008Corgosinho, P.H.C.; Martínez Arbizu, P. and Reid, J.W. 2008. Revision of the genus Murunducaris (Copepoda: Harpacticoida: Parastenocarididae), with descriptions of two new species from South America. Journal of Crustacean Biology, 28: 700-720.).
(3) Proximal row of spinules on the inner margin of the male P4 exp1. Males of Murunducaris and Santaremicaris gen. nov. have a proximal row of spinules on the inner margin of the P4 exp1. Within the Fontinalicaridinae, the Neotropical Forficatocaris, the African Pa. fossoris, Pa. crassicaudis (see Fischer, 1998Fischer, L. 1998. Parastenocarididae (Copepoda) aus Stränden des Malawisees (Ostafrika): Taxonomie, Vertikalverteilung und geographische Verbreitung. Oldenburg, Oldenburg University, Master’s Thesis. [Unpublished]), Pa. nigerianus, Pa. jeanninei, and Pa. kimi, the European Proserpinicaris phyllura (incertae sedis) (see Glatzel, 1991Glatzel, T. 1991. Neue morphologische Aspekte und die Copepodid-Stadien von Parastenocaris phyllura Kiefer (Copepoda, Harpacticoida). Zoologica Scripta, 20: 375-393.), and maybe Pa. aberrans have a proximal inner row of acuminate spinules in the P4 exp1. In the absence of a complete phylogenetic analysis of the Fontinalicaridinae, we consider this ornamentation as synapomorphic for Murunducaris and Santaremicaris gen. nov. only.
Characters separating Santaremicaris gen. nov. from Murunducaris
The new genus differs from Murunducaris by (4a) the presence of ornamentation on the inner margin of the P1 enp1 of male and female (Pl) (also present in M. juneae), (4b) such ornamentation is represented by stiff and long spinules (Ap); (5a) the absence of distal spinules on the enp of the male P2 (Ap), (5b) which is ornamented with strong outer spinules (Ap) and (5c) armed with long distal seta (Ap); (6) absence of a distal spine on the apophysis of male P3 (Ap); (7a) presence of a long row of spinules on the inner margin of the basis of male P3 (Ap), (7b) which has no trace of enp or seta indicating the enp (Ap); (8) presence of a triangular, short and sclerotized enp in the male P4 (Ap); (9) Presence of an inner distal row of spinules on the male P4 exp1; (10a) presence of a long and slim male P5 (Pl), (10b) with irregular margins (Ap), (10c) ornamented on the inner margin (Pl), (10d) ending in a long tip (probably representing a fused distal atrophied spine; Ap?), and (10e) with only three setigerous elements on the outer margin (Ap); (11) absence of an integumental window in the male second urosomite (Ap).
(4a−b) Presence of long and stiff spinules on the inner margin of the P1 enp1. The presence of an inner ornamentation on the enp1 of P1 is considered to be a plesiomorphic condition (Corgosinho et al., 2007bCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2007b. Three new species of Remaneicaris Jakobi, 1972 (Copepoda, Harpacticoida, Parastenocarididae) from the Ribeirão do Ouro River, Minas Gerais, Brazil, with some remarks on the ground pattern Parastenocarididae. Zootaxa, 1437: 1-28.). Within the Fontinalicaridinae genera, this ornamentation is observed in Fontinalicaris (see Martínez Arbizu, 1997Martínez Arbizu, P. 1997. Parastenocaris hispanica n. sp. (Copepoda: Harpacticoida: Parastenocarididae) from hyporheic groundwaters in Spain and its phylogenetic position within the fontinalis-group of species. Contributions to Zoology, 66: 215-226.; Schminke, 2010Schminke, H.K. 2010. High-level phylogenetic relationships within Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 83: 343-367.), Proserpinicaris (see Karanovic et al., 2012Karanovic, T.; Cho, J-L. and Lee, W. 2012. Redefinition of the parastenocaridid genus Proserpinicaris (Copepoda: Harpacticoida), with description of three new species from Korea. Journal of Natural History, 46: 25-26.; Karanovic and Lee, 2012Karanovic, T. and Lee, W. 2012. A new species of Parastenocaris from Korea, with a redescription of the closely related P. biwae from Japan (Copepoda: Harpacticoida: Parastenocarididae). Journal of Species Research, 1: 4-34.; Totakura et al., 2014Totakura, V.R.; Ranga-Reddy, Y. and Shaik, S. 2014. Three new species of Parastenocarididae (Crustacea, Copepoda, Harpacticoida) from India. Zootaxa, 3821: 501-537.), Brasilibathynellocaris (except in the male of Brasilibathynellocaris brasilibathynellae (Jakobi and Loyola e Silva, 1962Jakobi, H. and Loyola e Silva , J. 1962. Two new species of Parastenocaris (Copepoda: Harpacticoidea) from Santa Catarina, Brazil. Proceeding of the United States National Museum, 113: 389-397.) see Corgosinho et al., 2010aCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2010a. Revision of Brasilibathynellocaris Jakobi, 1972 (Copepoda: Harpacticoida: Parastenocarididae) with redefinition of the genus. Zoological Journal of the Linnean Society, 159: 527-566.), Iticocaris (see Corgosinho et al., 2012aCorgosinho, P.H.C.; Martínez Arbizu, P. and Previatelli, D. 2012a. Establishment of a new genus for Parastenocaris itica (Copepoda, Harpacticoida) from El Salvador, Central America, with discussion of the Parastenocaris fontinalis and P. proserpina groups. Iheringia, Série Zoologia, 102: 401-411. ), Siolicaris (except in Siolicaris sioli (Noodt, 1963Noodt, W. 1963. Subterrane Crustaceen der zentralen Neotropis. Zur Frage mariner Relikte im Bereich des Rio Paraguay - Paraná - Amazonas Systems. Zoologischer Anzeiger, 171: 114-147.), see Corgosinho et al., 2012bCorgosinho, P.H.C.; Ranga Reddy, Y. and Martínez Arbizu, P. 2012b. Revision of the genus Siolicaris Jakobi, 1972, with redescriptions of S. sioli (Noodt, 1963) and S. jakobi (Noodt, 1963) from South America, and S. sandhya (Ranga Reddy, 2001) comb. nov. from India (Copepoda, Harpacticoida, Parastenocarididae). Zootaxa, 3493: 49-71.), Colombocaris (see Gaviria et al., 2017Gaviria, S.; Defaye, D. and Corgosinho, P.H.C. 2017. Two new parastenocaridid genera (Copepoda, Harpacticoida) from South America, including a revision of Noodt’s columbiensis-group and the redescription of Parastenocaris tapajoensis Noodt, 1963. Crustaceana, 90: 535-570. ), Noodtcaris (see Noodt, 1972Noodt, W. 1972. Brasilianische Grundwasser-Crustacea, 1. Studien an den Gattungen Parastenocaris Kessler und Forficatocaris Jakobi aus der Serra do Mar von São Paulo (Copepoda, Harpacticoida). Crustaceana, 23: 76-99.; Gaviria et al., 2017Gaviria, S.; Defaye, D. and Corgosinho, P.H.C. 2017. Two new parastenocaridid genera (Copepoda, Harpacticoida) from South America, including a revision of Noodt’s columbiensis-group and the redescription of Parastenocaris tapajoensis Noodt, 1963. Crustaceana, 90: 535-570. ), and Santaremicaris gen. nov. Within Forficatocaris and Potamocaris one or more spinules are modified in the male, this being a synapomorphy shared by both genera; females retain the plesiomorphic condition. Within Murunducaris, it is absent in all species, except for M. juneae (see Corgosinho et al., 2008Corgosinho, P.H.C.; Martínez Arbizu, P. and Reid, J.W. 2008. Revision of the genus Murunducaris (Copepoda: Harpacticoida: Parastenocarididae), with descriptions of two new species from South America. Journal of Crustacean Biology, 28: 700-720.). The presence of a row of spinules on the inner margin of the P1 enp1 appears to be a widespread characteristic. This condition is present in Fontinalicaridinae and Parastenocaridinae (see Ranga Reddy et al., 2014Ranga Reddy, Y.; Totakura, V.R. and Corgosinho, P.H.C. 2014. Himalayacaris alaknanda n. gen., sp. nov. (Copepoda: Harpacticoida: Parastenocarididae) from the hyporheic zone of a Himalayan River, northern India. Journal of Crustacean Biology, 34: 801-819., fig. 9 and Corgosinho et al., 2017bCorgosinho, P.H.C.; Schizas, N.V.; Previattelli, D.; Rocha, C.E.F. da and Santos-Silva, E.N. 2017b. A new genus of Parastenocarididae (Copepoda, Harpacticoida) from the Tocantins River basin (Goiás, Brazil), and a phylogenetic analysis of the Parastenocaridinae. Zoosystematics and Evolution, 93: 167-187.). However, the spinules present in Santaremicaris gen. nov. are longer and more numerous than in other Neotropical species. Within Brasilibathynellocaris, stiff spinules are present only in one species, Brasilibathynellocaris paranaensisJakobi and Loyola e Silva, 1962Jakobi, H. and Loyola e Silva , J. 1962. Two new species of Parastenocaris (Copepoda: Harpacticoidea) from Santa Catarina, Brazil. Proceeding of the United States National Museum, 113: 389-397. (Corgosinho et al., 2010aCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2010a. Revision of Brasilibathynellocaris Jakobi, 1972 (Copepoda: Harpacticoida: Parastenocarididae) with redefinition of the genus. Zoological Journal of the Linnean Society, 159: 527-566.). In Noodtcaris, only N. tapajoensis (see Gaviria et al., 2017Gaviria, S.; Defaye, D. and Corgosinho, P.H.C. 2017. Two new parastenocaridid genera (Copepoda, Harpacticoida) from South America, including a revision of Noodt’s columbiensis-group and the redescription of Parastenocaris tapajoensis Noodt, 1963. Crustaceana, 90: 535-570. ) has this character. Therefore, we consider the presence of stiff spinules in M. juneae, B. paranaensis, and N. tapajoensis as convergent to the condition observed in Santaremicaris gen. nov.
(5a−c) Ornamentation and armature of male P2 enp. The condition found in S. santaremensis comb. nov. and S. amazonensis sp. nov. is unique within the family, not to mention within the Neotropical Fontinalicaridinae, and consists of a clear synapomorphy uniting both species. In males of Santaremicaris gen. nov. the distal seta is longer than usual, and its insertion occupies almost the whole distal margin of the enp. Additionally, the ornamentation is strong and limited to the outer margin only, with no distal or inner ornamentation. A P2 enp highly dimorphic in shape, also differing in ornamentation is not a widespread character within the Parastenocarididae. In most genera and species the P2 enp is very similar in female and male, differing in fine ornamentation only. The morphology, normally, is represented by an enp not too short or longer than half of the exp1, with a small distal seta, with an insertion not as wide as the distal margin of the segment, with some few distal spinules and some few outer and maybe inner spinules. Within the Parastenocaridinae it is reduced to a small enp with a distal seta in Monodicaris, but is normally developed in most species within this subfamily. Within the Fontinalicaridinae it is long and strongly ornamented in males of B. brasilibathynellae and B. paranaensis, occurring as a derived condition within this genus (Corgosinho et al., 2010aCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2010a. Revision of Brasilibathynellocaris Jakobi, 1972 (Copepoda: Harpacticoida: Parastenocarididae) with redefinition of the genus. Zoological Journal of the Linnean Society, 159: 527-566.). It is claviform, strongly ornamented, with a distal seta and accompanied by a distal spinule bearing a hyaline margin in S. sandhya, but normally developed in the remaining Siolicaris species described so far. All the species of Murunducaris have some dimorphism in the P2 enp. A foliaceous P2 enp is more developed in M. juneae, M. loyolai, and M. noodti. In M. dactyloides it is less conspicuous. A similar kind of sexual dimorphism can be observed in Iticocaris itica (Noodt, 1962Noodt, W. 1962. Limnisch-subterrane Copepoden der Gattung Parastenocaris Kessler aus Mittelamerika. Beiträge zur Neotropischen Fauna, 2(3): 223-248.), Pa. fossoris, and Pa. crassicaudis, and in species closely related to Fontinalicaris fontinalis (Schnitter and Chappuis, 1915Schnitter, H. and Chappuis, P.A. 1915. Parastenocaris fontinalis nov. spec., ein neuer Süsswasserharpacticide. Zugleich ein Beitrag zur Kenntnis der Gattung Parastenocaris Kessler. Zoologischer Anzeiger, 45(7): 290-302.) (e.g., Fontinalicaris hispanicaMartínez Arbizu, 1997Martínez Arbizu, P. 1997. Parastenocaris hispanica n. sp. (Copepoda: Harpacticoida: Parastenocarididae) from hyporheic groundwaters in Spain and its phylogenetic position within the fontinalis-group of species. Contributions to Zoology, 66: 215-226.). A similar condition seems to be present in Noodtcaris, but this is difficult to evaluate based only on the original descriptions. Corgosinho et al. (2008Corgosinho, P.H.C.; Martínez Arbizu, P. and Reid, J.W. 2008. Revision of the genus Murunducaris (Copepoda: Harpacticoida: Parastenocarididae), with descriptions of two new species from South America. Journal of Crustacean Biology, 28: 700-720.) mentioned that a dimorphism of the P2 enp is absent in Forficatocaris, but the study of new and undescribed species (unpublished data), and the reexamination of the literature and some type material revealed that the P2 enp is thick and/or long with some few minute spinules along the inner margin of the distal 1/3 in some males of Forficatocaris (e.g., Forficatocaris evelinae Noodt, 1972; Forficatocaris lilianae Noodt, 1972; Forficatocaris odeteaeMoreira, Eskinazi-Sant’Anna and Previattelli, 2019Moreira, R.A.; Sant’Anna, E.M.R. and Previattelli, D. 2019. Forficatocaris odeteae n. sp., a new parastenocarididae (copepoda, harpacticoida) from a high-altitude pond in Minas Gerais State, Brazil. Crustaceana, 92: 163-176.).
(6) Male P3 apophysis lacking distal spine. According to Schminke (2010Schminke, H.K. 2010. High-level phylogenetic relationships within Parastenocarididae (Copepoda, Harpacticoida). Crustaceana, 83: 343-367.), there is no distal spine on the apophysis of a Parastenocaridinae. The presence of this armature in Fontinalicaridinae and its absence in Parastenocaridinae is considered a plesiomorphic and an apomorphic condition, respectively (Corgosinho et al., 2017bCorgosinho, P.H.C.; Schizas, N.V.; Previattelli, D.; Rocha, C.E.F. da and Santos-Silva, E.N. 2017b. A new genus of Parastenocarididae (Copepoda, Harpacticoida) from the Tocantins River basin (Goiás, Brazil), and a phylogenetic analysis of the Parastenocaridinae. Zoosystematics and Evolution, 93: 167-187.). Within the Fontinalicaridinae, an apophysis with a distal spine occurs in Fontinalicaris, Proserpinicaris partim, Murunducaris, some species of Forficatocaris and Potamocaris, S. sandhya and Siolicaris xyrophora (Noodt and Galhano, 1969Noodt, W. and Galhano, M.H. 1969. Studien an Crustacea Subterranea (Isopoda, Syncarida, Copepoda) aus dem Norden Portugals. Publicações do Instituto de Zoologia ‘Dr. Augusto Nobre’, 107:1-75.), N. tapajoensis and N. kubitzkii, and other fontinalicaridids still included in the genus Parastenocaris, such as Pa. fossoris, and perhaps in the Portuguese Proserpinicaris cruzi (Noodt and Galhano, 1969Noodt, W. and Galhano, M.H. 1969. Studien an Crustacea Subterranea (Isopoda, Syncarida, Copepoda) aus dem Norden Portugals. Publicações do Instituto de Zoologia ‘Dr. Augusto Nobre’, 107:1-75.) (incertae sedis). In all these cases, except for S. sandhya and S. xyrophora, which are derived species within Siolicaris, the armature is considered the retention of a plesiomorphic character. The apophysis with a clear incorporated spine occurs in the Colombian N. columbiensis, Noodtcaris roettgeri (Noodt, 1972Noodt, W. 1972. Brasilianische Grundwasser-Crustacea, 1. Studien an den Gattungen Parastenocaris Kessler und Forficatocaris Jakobi aus der Serra do Mar von São Paulo (Copepoda, Harpacticoida). Crustaceana, 23: 76-99.), and in some Proserpinicaris species. The spine is completely absent in the Neotropical Brasilibathynellocaris, Siolicaris, Iticocaris itica, some Potamocaris species, Santaremicaris gen. nov., Parastenocaris staheliMenzel, 1916Menzel, R. 1916. Über das Auftreten der Harpacticiden gattungen Epactophanes Mrazek und Parastenocaris Kessler in Surinam. Zoologischer Anzeiger, 47: 145-152., the African Pa. crassicaudis, Pa. kimi, Pa. nigerianus, and Pa. jeanninei, the Spanish Proserpinicaris cantabrica (Chappuis, 1937Chappuis, P.A. 1937. Subterrane Harpacticoiden aus Nord-Spanien. Buletinul Societatii de Stiinte din Cluj, 8: 556-571.) (incertae sedis), Parastenocaris calliroeCottarelli, 1969Cottarelli, V. 1969. Una nuova Parastenocaris di Sardegna (Crustacea, Copepoda, Harpacticoida). Rendiconti dell’Istituto Lombardo di Scienze e Lettere (B), 103: 255-268. (Sardinia), and Parastenocaris sardoaCottarelli and Torrisi, 1977Cottarelli, V. and Torrisi, M.R. 1977. Su alcune Parastenocaris di acque interstiziali sarde (Crustacea, Copepoda, Harpacticoida). Fragmenta Entomologica, Rome, 13: 285-297. (Sardinia). This is a heterogeneous group of genera and species probably belonging to different evolutionary lineages, some of them not directly related to each other. Hence, it is more parsimonious to consider the loss of the distal spine occurring several times within the Fontinalicaridinae. Within the Neotropical zone, the loss of the distal spine of the male P3 apophysis would have occurred independently within the monophylum Potamocaris/Forficatocaris, because the presence of the distal spine is in the ground pattern of this clade. It occurs again in the putative monophylum Brasilibathynellocaris/Iticocaris/Siolicaris, and once in the lineage of Santaremicaris gen. nov. within the monophylum Murunducaris/Santaremicaris gen. nov.
(7a−b) male P3 basis with a long row of spinules on the inner margin and enp absent. Within the Fontinalicaridinae an inner row of spinules on the basis of the male P3 is present in N. tapajoensis, Pa. crassicaudis, Pa. fossoris, Pa. nigerianus, Pa. kimi, Pa. sardoa, Pa. calliroe, Proserpinicaris phyllura (incertae sedis), and some Proserpinicaris (e.g., Proserpinicaris admete (Cottarelli, Fasano, Mura and Saporito, 1980Cottarelli, V.; Fasano, L.; Mura, G. and Saporito P.E. 1980. Parastenocaris adméte n. sp. (Crustacea, Copepoda, Harpacticoida) di acque interstiziali di Sardegna. Rivista di Idrobiologia, 19:619-628); Proserpinicaris ima (Cottarelli, 1989Cottarelli, V. 1989. Ricerche zoologiche della nave oceanografica ‘Minerva’ (C.N.R.) sulle isole circumsarde. IV. Un nuovo arpacticoide (Crustacea, Copepoda) freatobio dell’isola La Maddalena (Sardegna): Parastenocaris ima n. sp. Annali del Museo Civico de Storia Naturale ‘Giacomo Doria’, Genova, 87: 285-296.), Proserpinicaris kalypso (Pesce, Galassi and Cottarelli, 1988Pesce, G.L.; Galassi, D.M.P. and Cottarelli, V. 1988. First representative of the family Parastenocaridae from Sicily (Italy), and description of two new species of Parastenocaris Kessler (Crustacea Copepoda:Harpacticoida). Bulletin Zoölogisch Museum, Universiteit van Amsterdam, 11: 137-141.); Proserpinicaris proserpina (Chappuis, 1938Chappuis, P.A. 1938. Subterrane Harpacticoiden aus Süd-Italien. Buletinul Societatii de Stiinte din Cluj, 9: 153-181.)), S. xyrophora, and maybe in Pr. cruzi (incertae sedis). Within Potamocaris and Forficatocaris the inner row of spinules is fused into a process (“Vorsprung”), which is a synapomorphy uniting these genera. Inner basal ornamentation of the male P3 is absent in Colombocaris, Iticocaris, Murunducaris, Noodtcaris, Siolicaris, Pa. staheli, Fontinalicaris sensuMartínez Arbizu (1997Martínez Arbizu, P. 1997. Parastenocaris hispanica n. sp. (Copepoda: Harpacticoida: Parastenocarididae) from hyporheic groundwaters in Spain and its phylogenetic position within the fontinalis-group of species. Contributions to Zoology, 66: 215-226.), Pa. jeanninei, Parastenocaris matopoicaWells, 1964Wells, J.B.J. 1964. Six new species of Parastenocaris (Crustacea, Copepoda) from southern Rhodesia. Annals and Magazine of Natural History, 7(13): 193-204., Pa. aberrans, Pr. cantabrica (incertae sedis), and other Proserpinicaris (e.g., Proserpinicaris corgosinhoi Totakura, Ranga-Reddy and Shaik, 2014Totakura, V.R.; Ranga-Reddy, Y. and Shaik, S. 2014. Three new species of Parastenocarididae (Crustacea, Copepoda, Harpacticoida) from India. Zootaxa, 3821: 501-537.; Proserpinicaris imjinKaranovic, Cho and Lee, 2012Karanovic, T.; Cho, J-L. and Lee, W. 2012. Redefinition of the parastenocaridid genus Proserpinicaris (Copepoda: Harpacticoida), with description of three new species from Korea. Journal of Natural History, 46: 25-26.; Proserpinicaris moravica (Sterba, 1965Sterba, O. 1965. Ueber eine neue Parastenocaris-Art (Copepoda-Harpacticoidea) aus Mähren. Zoologischer Anzeiger, 175(3): 204-208.); Proserpinicaris nipponensis (Chappuis, 1955Chappuis, P.A. 1955. Harpacticoïdes psammiques du lac Tanganika. Revue de Zoologie et de Botanique Africaines, 51: 68-80.); Proserpinicaris ondali (Lee and Chang, 2009Lee, J.M. and Chang, C.Y. 2009. Two Groundwater Copepods of the Genus Parastenocaris (Harpacticoida, Parastenocarididae) from South Korea. Animal Cells and Systems, 13: 169-178.); Proserpinicaris wangpiKaranovic, Cho and Lee, 2012Karanovic, T.; Cho, J-L. and Lee, W. 2012. Redefinition of the parastenocaridid genus Proserpinicaris (Copepoda: Harpacticoida), with description of three new species from Korea. Journal of Natural History, 46: 25-26.; Proserpinicaris youngKaranovic, Cho and Lee, 2012Karanovic, T.; Cho, J-L. and Lee, W. 2012. Redefinition of the parastenocaridid genus Proserpinicaris (Copepoda: Harpacticoida), with description of three new species from Korea. Journal of Natural History, 46: 25-26.). The male P3 enp is not illustrated or it is absent in Lacustricaris budapestiensis (Török, 1935Török, P. 1935. A Budapesti Vízvezetéki víz szüredékének faunája. Matematikai es Természettudomanyi Ertesito, Budapest, 53: 637-664.) Parastenocaris lacustrisChappuis, 1958Chappuis, P.A. and Delamare Deboutteville, C. 1958. Recherches sur la faune interstitielle littorale du Lac Érié. Le problème des glaciations quaternaires. Vie et Milieu 8: 366-376. (incertae sedis) (Chappuis and Delamare Deboutteville, 1958Chappuis, P.A. and Delamare Deboutteville, C. 1958. Recherches sur la faune interstitielle littorale du Lac Érié. Le problème des glaciations quaternaires. Vie et Milieu 8: 366-376.), Pa. aberrans, Pa. nigerianus, Pa. matopoica, Pa. jeanninei, Pa. staheli and some Proserpinicaris.
It seems that an inner basal ornamentation evolved independently within the Fontinalicaridinae and its presence is not necessarily connected to the presence of the enp in some species such as Pa. calliroe, Pa. nigerianus, and at least Pr. ima within Proserpinicaris. Neither the absence of ornamentation is associated with the absence of the enp in all cases. However, the enp is sometimes represented by a small spine or seta, which makes its identification difficult and it was possibly omitted in some old descriptions. The absence of both enp and ornamentation undisputedly occurs in Colombocaris, Iticocaris, M. juneae and M. dactyloides.
In the ground pattern of Murunducaris the male P3 basis is represented by an unadorned segment with a spiniform enp. Instead, a basis without an enp and with a long row of spinules on the inner margin is present only in the ground pattern of Santaremicaris gen. nov. We believe that the condition present in Santaremicaris gen. nov. is homologous to what we observed in Pa. fossoris, Pa. crassicaudis, Pa. kimi, and Pa. nigerianus. In these species, the ornamentation does not occupy the whole inner surface of the basis and the enp is posteriorly inserted to it. We hypothesize that this is a plesiomorphic condition. In Santaremicaris gen. nov. the basis of the male P3 is elongated, ornamented with a long row of minute spinules along the inner margin, and the enp is absent; this being considered an autapomorphy supporting this genus.
(8) Triangular, short and sclerotized enp of the male P4. A short and sclerotized enp of the male P4 is a unique feature of Santaremicaris gen. nov. No other Fontinalicaridinae has these combined features. A superficially similar enp is observed in Pr. nipponensisChappuis, 1955Chappuis, P.A. 1955. Harpacticoïdes psammiques du lac Tanganika. Revue de Zoologie et de Botanique Africaines, 51: 68-80., and Pa. nigerianus. However, based on the illustrations of these species, we can say nothing about sclerotization. Sclerotized male P4 enp appear in the monophylum Potamocaris/Forficatocaris, especially in Forficatocaris. However, the morphology of the P4 enp in Forficatocaris is so different from that of Santaremicaris gen. nov. that any hypothesis of homology must be abandoned. The male P4 enp in the remaining fontinalicaridids have a variety of shapes, i.e., long and lamelliform, foliaceous and hirsute, spiniform, short and cuneiform, or even absent. In Murunducaris, the sister group of Santaremicaris gen. nov., the male P4 enp is long, lamelliform with outer ornamentation as in M. dactyloides and M. noodti or slightly bent inwards, with ornamentation mostly distributed outwardly as in M. juneae and M. loyolai. However, in no Murunducaris species is the enp sclerotized. Therefore, we consider the triangular, short and sclerotized enp of the male P4 as autapomorphic for Santaremicaris gen. nov.
(9) Presence of an inner distal row of spinules on the male P4 exp1. During the ontogeny of most Parastenocarididae, a distal inner row of spinules on the male P4, still present in the copepodid V (see Glatzel, 1991Glatzel, T. 1991. Neue morphologische Aspekte und die Copepodid-Stadien von Parastenocaris phyllura Kiefer (Copepoda, Harpacticoida). Zoologica Scripta, 20: 375-393.; Corgosinho et al., 2010aCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2010a. Revision of Brasilibathynellocaris Jakobi, 1972 (Copepoda: Harpacticoida: Parastenocarididae) with redefinition of the genus. Zoological Journal of the Linnean Society, 159: 527-566.) is homologous to the adult hyaline frill. The presence of the inner hyaline frill in the adults is considered a plesiomorphic condition of the Parastenocarididae, and the retention of the juvenile character in the adult (a distal inner row of spinules) is the apomorphic condition.
A distal inner row of spinules is present in S. santaremensis comb. nov. and in S. amazonensis sp. nov. The same character is also observed in Forficatocaris, Pa. fossoris, Pa. crassicaudis, Pa. jeanninei, Pa. kimi, Pa. nigerianus, Pr. phyllura (incertae sedis), Fontinalicaris (Parastenocaris fontinalis-group sensuMartínez Arbizu, 1997Martínez Arbizu, P. 1997. Parastenocaris hispanica n. sp. (Copepoda: Harpacticoida: Parastenocarididae) from hyporheic groundwaters in Spain and its phylogenetic position within the fontinalis-group of species. Contributions to Zoology, 66: 215-226.; Fig. 5A, B), and Proserpinicaris.
(10a−e) Shape, ornamentation, and armature of the male P5. The shared presence of an enlarged male P5 intercoxal sclerite with a cuticular process between the limbs in both Santaremicaris gen. nov. and Murunducaris justify a sister group relationship between these genera. An enlarged intercoxal sclerite was reported by Corgosinho et al. (2008Corgosinho, P.H.C.; Martínez Arbizu, P. and Reid, J.W. 2008. Revision of the genus Murunducaris (Copepoda: Harpacticoida: Parastenocarididae), with descriptions of two new species from South America. Journal of Crustacean Biology, 28: 700-720.) for Pa. fossoris and Pa. crassicaudis, but this condition is not illustrated by Fischer (1998Fischer, L. 1998. Parastenocarididae (Copepoda) aus Stränden des Malawisees (Ostafrika): Taxonomie, Vertikalverteilung und geographische Verbreitung. Oldenburg, Oldenburg University, Master’s Thesis. [Unpublished]). The same condition may be present in Pa. aberrans from the Northern Pyrenees in France (Apostolov, 2004Apostolov, A. 2004. Copépodes harpacticoïdes stygobies de France. 6. Description d’un nouveau représentant du genre Parastenocaris Kessler, 1913. Crustaceana, 77: 197-203.).
One of the most striking characters of the genus Murunducaris is the presence of a large, subconical, terminal and unfused spine on the P5 of males (Corgosinho et al., 2008Corgosinho, P.H.C.; Martínez Arbizu, P. and Reid, J.W. 2008. Revision of the genus Murunducaris (Copepoda: Harpacticoida: Parastenocarididae), with descriptions of two new species from South America. Journal of Crustacean Biology, 28: 700-720.). This transformation is unique for Murunducaris. However, the observation of the P5 of S. santaremensis comb. nov., S. amazonensis sp. nov., Pa. crassicaudis, Pa. fossoris, Pa. aberrans, and Pa. jeanninei revealed very interesting conditions. In Pa. fossoris, the distal margin of the P5 is represented by a much-ornamented triangular region (Fischer, 1998Fischer, L. 1998. Parastenocarididae (Copepoda) aus Stränden des Malawisees (Ostafrika): Taxonomie, Vertikalverteilung und geographische Verbreitung. Oldenburg, Oldenburg University, Master’s Thesis. [Unpublished]; Fig. 11A), and on the outer margin, there are only two setae between this ornamented structure and the articulated setae of the basis. A similar P5 of male, but without the distal ornamentation, is described for Pa. matopoica. In Pa. crassicaudis (see Fischer, 1998Fischer, L. 1998. Parastenocarididae (Copepoda) aus Stränden des Malawisees (Ostafrika): Taxonomie, Vertikalverteilung und geographische Verbreitung. Oldenburg, Oldenburg University, Master’s Thesis. [Unpublished]; Fig. 11B) and Pa. jeanninei, a strong spine appears on the distal margin of P5, but a clear suture indicates the point of insertion of this spine. In Pa. aberrans, the distal spine may be what Apostolov (2004Apostolov, A. 2004. Copépodes harpacticoïdes stygobies de France. 6. Description d’un nouveau représentant du genre Parastenocaris Kessler, 1913. Crustaceana, 77: 197-203.) depicts as an inner hyaline lamella. In S. santaremensis comb. nov. and S. amazonensis sp. nov., there are only three setae on the outer margin and a distal spiniform process. This distal spiniform process may be a fused distal spine, homologous to the distal ornamented process of Pa. fossoris. We consider that the unfused condition of the distal spine of Murunducaris, Pa. crassicaudis, Pa. jeanninei, and Pa. aberrans is a plesiomorphy. Autapomorphies of Santaremicaris gen. nov. are long male P5 with irregular margins and with a putative fused distal spine.
Parastenocaris fossoris, male P5 (A). Parastenocaris crassicaudis, male P5 (B). Scale bars 50µm. After Fischer (1998Fischer, L. 1998. Parastenocarididae (Copepoda) aus Stränden des Malawisees (Ostafrika): Taxonomie, Vertikalverteilung und geographische Verbreitung. Oldenburg, Oldenburg University, Master’s Thesis. [Unpublished]). Reproduction authorized by Dr. Lutz Fischer and Dr. Thomas Glatzel.
(11) Male second Ur without integumental window. It is very difficult to discuss the phylogenetic significance of the urosomal integumental windows on the Urs of Parastenocarididae. In most cases they are not described or are considered absent. Recently its phylogenetic and taxonomic importance was highlighted by Corgosinho and Martínez Arbizu (2005Corgosinho, P.H.C. and Martínez Arbizu, P. 2005. Two new interstitial species of Remaneicaris Jakobi (Copepoda, Harpacticoida, Parastenocarididae) from the Ribeirão do Ouro river, Brazil, with a redefinition of the genus. Senckenbergiana biologica, 85: 147-162.) and Corgosinho et al. (2007aCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2007a. Redescription of Remaneicaris ignotus (Dussart, 1983) a Parastenocarididae (Copepoda, Harpacticoida) with an unusual set of plesiomorphic characters. Invertebrate Zoology, 4: 31-44.; 2007bCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2007b. Three new species of Remaneicaris Jakobi, 1972 (Copepoda, Harpacticoida, Parastenocarididae) from the Ribeirão do Ouro River, Minas Gerais, Brazil, with some remarks on the ground pattern Parastenocarididae. Zootaxa, 1437: 1-28.) and Schminke (2008Schminke, H.K. 2008. First report of groundwater fauna from Papua New Guinea: Kinnecaris Jakobi, 1972 redefined (Copepoda, Harpacticoida, Parastenocarididae), and description of a new species. Crustaceana, 81: 1241-1253.; 2009Schminke, H.K. 2009. Monodicaris gen. n. (Copepoda, Harpacticoida, Parastenocarididae) from west Africa. Crustaceana, 82: 367-378). The most plesiomorphic pattern is the presence of dorsal integumental windows from Ur2 to Ur5 in male and Ur2 to Ur4 in females. This is considered the ground pattern for the Parastenocarididae by Corgosinho et al. (2007aCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2007a. Redescription of Remaneicaris ignotus (Dussart, 1983) a Parastenocarididae (Copepoda, Harpacticoida) with an unusual set of plesiomorphic characters. Invertebrate Zoology, 4: 31-44.). To our knowledge, lateral integumental windows have evolved only within the Parastenocaridinae, in Remaneicaris (Corgosinho and Martínez Arbizu, 2005Corgosinho, P.H.C. and Martínez Arbizu, P. 2005. Two new interstitial species of Remaneicaris Jakobi (Copepoda, Harpacticoida, Parastenocarididae) from the Ribeirão do Ouro river, Brazil, with a redefinition of the genus. Senckenbergiana biologica, 85: 147-162. and Corgosinho et al., 2007bCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2007b. Three new species of Remaneicaris Jakobi, 1972 (Copepoda, Harpacticoida, Parastenocarididae) from the Ribeirão do Ouro River, Minas Gerais, Brazil, with some remarks on the ground pattern Parastenocarididae. Zootaxa, 1437: 1-28., 2010bCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2010b. Three new interstitial species of Remaneicaris Jakobi, 1972 (Copepoda: Harpacticoida: Parastenocarididae) from southern and southeastern Brazil. Invertebrate Zoology, 7: 1-28., 2017aCorgosinho, P.H.C.; Mercado-Salas, N.; Martínez Arbizu, P.; Santos Silva, E.N. and Kihara, T.C. 2017a. Revision of the Remaneicaris argentina-group (Copepoda, Harpacticoida, Parastenocarididae): supplementary description of species, and description of the first semi-terrestrial Remaneicaris from the tropical forest of Southeast Mexico. Zootaxa, 4238: 499-530.), Kinnecaris (Schminke, 2008Schminke, H.K. 2008. First report of groundwater fauna from Papua New Guinea: Kinnecaris Jakobi, 1972 redefined (Copepoda, Harpacticoida, Parastenocarididae), and description of a new species. Crustaceana, 81: 1241-1253.; Bruno and Cottarelli, 2015Bruno, M.C. and Cottarelli, V. 2015. First record of Kinnecaris (Copepoda: Harpacticoida: Parastenocarididae) from Turkey and Thailand; description of three new species and emended definition of the genus. Italian Journal of Zoology, 82: 69-94.), and Monodicaris. Therefore, the ground pattern of Parastenocarididae is retained in Fontinalicaridinae. In the absence of contrary evidence, the loss of the integumental window on the Ur2 segment in male is considered as an autapomorphy of Santaremicaris gen. nov.
The curved hook on the inner margin of the basis of male P3.
A curved hook on the inner margin of the basis of male P3 could be considered a modified enp in Santaremicaris gen. nov. With exception of Remaneicaris (see Corgosinho et al., 2007bCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2007b. Three new species of Remaneicaris Jakobi, 1972 (Copepoda, Harpacticoida, Parastenocarididae) from the Ribeirão do Ouro River, Minas Gerais, Brazil, with some remarks on the ground pattern Parastenocarididae. Zootaxa, 1437: 1-28.), in which the enp is one-segmented, in the remaining Parastenocarididae the enp, when present, is represented by a seta or spine. According to Corgosinho et al. (2010aCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2010a. Revision of Brasilibathynellocaris Jakobi, 1972 (Copepoda: Harpacticoida: Parastenocarididae) with redefinition of the genus. Zoological Journal of the Linnean Society, 159: 527-566.), with the exception of B. paranaensis, all Brasilibathynellocaris species have a curved hook proximally inserted on the inner margin of male P3, similar to what is observed in S. santaremensis comb. nov. Within Brasilibathynellocaris, the curved hook is proximal to the seta representing the enp (Corgosinho et al., 2010aCorgosinho, P.H.C.; Martínez Arbizu, P. and Santos-Silva, E.N. 2010a. Revision of Brasilibathynellocaris Jakobi, 1972 (Copepoda: Harpacticoida: Parastenocarididae) with redefinition of the genus. Zoological Journal of the Linnean Society, 159: 527-566.). In addition, both in Santaremicaris gen. nov. and Brasilibathynellocaris the hook-like structure is not hollow, which is evidence that the curved hook is a spinule, instead of a modified spine. This character could be present in the common ancestor of Brasilibathynellocaris, Murunducaris, and Santaremicaris gen. nov. Brasilibathynellocaris could have retained the ancestral character (secondarily lost in B. paranaensis), whereas it was lost in the Murunducaris/Santaremicaris gen. nov. lineages, reappearing in S. santaremensis comb. nov. The expression of this character in S. santaremensis comb. nov. could be the result of the reactivation of a silent gene. Therefore, considering all the characters discussed in this contribution, it is difficult to discuss the evolution of the curved hook on the inner margin of the basis of male P3 without a proper phylogenetic analysis. We cannot exclude the possibility of convergent evolution.
Santaremicaris gen. nov. has a rectangular basis in the male P3 (two times longer than wide). A similar basis is described for Iticocaris and Brasilibathynellocaris, whereas Siolicaris, Potamocaris, Forficatocaris, and Murunducaris have a shorter basis. Despite the similarities mentioned above between Brasilibathynellocaris, Iticocaris and Santaremicaris gen. nov., to date, there is no strong evidence supporting a close affinity between these genera. Instead, the unique morphology of the P5 intercoxal sclerite and the armature of the P5 itself points to another direction.
Distinguishing characters between S. santaremensis comb. nov. and S. amazonensis sp. nov.
Santaremicaris santaremensis comb. nov. and S. amazonensis sp. nov. differ in body length, and the following male characters: a) ornamentation of the basis of P1; b) fine ornamentation of the enp1 of P1; c) fine ornamentation of the P2 enp; d) ornamentation of the coxa and basis of P3; e) shape and ornamentation of the P3 exp1; f) shape and ornamentation of P3 apophysis; g) length and shape of P3 thumb; h) fine ornamentation of P4 enp and exp1; i) fine ornamentation of the P4 exp3; j) shape of the intercoxal process on P5; and k) finally shape and fine ornamentation of P5. For the conditions of each character, see Tab. 1.
FINAL CONSIDERATIONS
It is still premature to say which group of species is closely related to Murunducaris and Santaremicaris gen. nov. However, as discussed above, there are some shared characters such as the presence of a proximal and distal row of spinules on the inner margin of male P4 exp1 (the latter eventually lost in some Murunducaris, Potamocaris, and Pa. aberrans) that could support a close relationship between Potamocaris, Forficatocaris, Murunducaris, Santaremicaris gen. nov., Pa. fossoris, Pa. crassicaudis, Pa. nigerianus, Pa. jeanninei, Pa. kimi, Pa. aberrans, and maybe Pr. phyllura (incertae sedis).
ACKNOWLEDGEMENTS
PHCC would like to thank the Deutscher Akademischer Austausch Dienst “DAAD”, the Fundação de Amparo à Pesquisa do Estado do Amazonas and the Forschungsinstitut Senckenberg for providing financial support. The work of PHCC at the University of Montes Claros, PPG-BURN, was supported by FAPEMIG (CRA−BPV−00393−16 and CRA−BPV−00547−17). PHCC and CEFR were financed by CNPq, grant number 563318/2010−4 and FAPESP, grant number 2010/52318−6 (project "Biodiversity of Microcrustaceans in Brazilian Rocky Fields"- SISBIOTA). We are indebted to the DZMB Senckenberg Research Institute and the Instituto Nacional de Pesquisas da Amazônia for the logistic support during this work; Prof. Dr. H.K. Schminke (University of Oldenburg) for discussions on parastenocaridid phylogeny; and Dr. Thomas Glatzel (University of Oldenburg) for allowing us to study his personal collection of Parastenocarididae. We are also thankful to Dr. Lutz Fischer and Dr. Thomas Glatzel who allowed the use of two illustrations in this work. We are thankful to Dr. H.W. Mittmann, director of the Kiefer's collection at the Staatliches Museum für Naturkunde Karlsruhe (Germany). This study would not have been possible without the study of Noodt´s type material. We express our special gratitude to Dr. Ahmed Ahnert who curated Noodt´s material and put it at our disposal for the present study at the DZMB.
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- Walter, T.C. and Boxshall, G. 2020. World of Copepods database. Parastenocarididae Chappuis, 1940. Accessed through: World Register of Marine Species at: Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=taxdetails&id=115170 Acessed on 05 november 2020.
» http://www.marinespecies.org/aphia.php?p=taxdetails&id=115170 - Wells, J.B.J. 1964. Six new species of Parastenocaris (Crustacea, Copepoda) from southern Rhodesia. Annals and Magazine of Natural History, 7(13): 193-204.
- Willen, E. 2000. Phylogeny of the Thalestridimorpha Lang, 1944 (Crustacea, Copepoda). Cuvillier Verlag, Göttingen . 233p.
Data availability
Data citations
Walter, T.C. and Boxshall, G. 2020. World of Copepods database. Parastenocarididae Chappuis, 1940. Accessed through: World Register of Marine Species at: Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=taxdetails&id=115170 Acessed on 05 november 2020.
Publication Dates
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Publication in this collection
14 May 2021 -
Date of issue
2021
History
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Received
02 Sept 2020 -
Accepted
25 Nov 2020