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ExopodEndopod
P20.0.0210.110
P30.0.0210.010
P40.0.1210.020
+ + +P3 endopod +♂ +not sexually dimorphic, two-segmented. P5 with outer basal seta and four discrete elements; innermost element fused to segment forming spinous process; length of process sometimes sexually dimorphic. P +6 ♂ +with two setae/spines. + + + + + +Type +species. + + +Arenopontia +( +Neoleptastacus +) +speluncae +Cottarelli, Bruno & Venanzetti, 1994 + +[by original designation]. + + +Other species. + +Arenopontia +( +Neoleptastacus +) +phreatica +Cottarelli, Bruno & Venanzetti, 1994 + += + +Phreatipontia phreatica +( +Cottarelli, Bruno & Venanzetti, 1994 +) + + +comb. nov. + + + + + +Etymology. +The genus name is derived from the Greek φρέαρ (phréar), meaning well, spring, and πόντος (pόntos), meaning sea, and refers to the low salinity habitat preference of its members. Gender: feminine. + + + + +Remarks. +The two members included in this genus, + +P. phreatica + + +comb. nov. + +and + +P. speluncae + + +comb. nov. + +, differ from all + +Neoleptastacus +species + +in the presence of two geniculate setae on the distal segment of the P1 endopod (instead of an outer spine and an inner geniculate seta). This character state is shared with two other genera in the +Arenopontiidae +, + +Psammoleptastacus + +and + +Onychopontia + +, both of which display sexual dimorphism on the P3 endopod which is not expressed in + +Phreatipontia + + +gen. nov. + + +Psammoleptastacus + +additionally differs from the new genus in (1) the presence of a dorsolateral spur on the inner margin of the caudal ramus, (2) P1 endopod being distinctly shorter than the exopod, (3) the presence of two distal elements on P2–P3 enp-2, and (4) the absence of an inner spinous process on the P +5 in +both sexes. + +Onychopontia + +can be differentiated from + +Phreatipontia + + +gen. nov. + +by (1) the characteristic deeply incised hyaline frills on the abdominal somites, (2) the absence of the inner serrate seta on P2 enp-2, (3) the very short P2–P3 enp-2, and (4) the absence of an inner spinous process on the female P5. Both species of + +Phreatipontia + + +gen. nov. + +display a reduced armature on P2–P3 endopods with only one apical seta on the distal segment. This condition is shared with members of the + +trisetosus + +-group of + +Neoleptastacus + +, however in this lineage the endopods are only 1-segmented ( +Table 2 +). Finally, both + +P. phreatica + + +comb. nov. + +and + +P. speluncae + + +comb. nov. + +exhibit a characteristic caudal ramus seta IV which has a long setule on the outer margin (indicated by arrows in +Figs 12C +; +16C +) which can be considered an autapomorphy for the genus. Current records suggest that, unlike most arenopontiids, both species favour low salinity environments, including phreatic and water table habitats ( + +Cottarelli +et al. +1994 + +, +1996 +; + +Bruno +et al. +1998 + +; this study). + + +Within the family, only + +Arenopontia +cf. +subterranea + +was previously recorded from habitats with a strong freshwater influence in various localities in Abruzzi, +Lazio +and +Tuscany +in +Italy +( +Cottarelli 1969 +; + +Cottarelli +et al. +1994 + +; +Cottarelli & Venanzetti 1989 +), representing a second but independent incursion into low salinity environments. + + + + \ No newline at end of file diff --git a/data/62/7E/C6/627EC678F74AFF92FF4EFE577DC2FF76.xml b/data/62/7E/C6/627EC678F74AFF93FF4EFE5778EEF8EF.xml similarity index 83% rename from data/62/7E/C6/627EC678F74AFF92FF4EFE577DC2FF76.xml rename to data/62/7E/C6/627EC678F74AFF93FF4EFE5778EEF8EF.xml index 882a7379652..0de953ab83d 100644 --- a/data/62/7E/C6/627EC678F74AFF92FF4EFE577DC2FF76.xml +++ b/data/62/7E/C6/627EC678F74AFF93FF4EFE5778EEF8EF.xml @@ -1,67 +1,67 @@ - - - -Review of Neoleptastacus Nicholls, 1945 (Harpacticoida, Arenopontiidae), including an updated key to species and proposal of Phreatipontia gen. nov. + + + +Review of Neoleptastacus Nicholls, 1945 (Harpacticoida, Arenopontiidae), including an updated key to species and proposal of Phreatipontia gen. nov. - - -Author + + +Author -Sak, Serdar -0000-0002-4955-3162 -Department of Biology, Faculty of Science and Literature, Balıkesir University, Balıkesir, Türkiye. serdarsak @ gmail. com. https: // orcid. org / 0000 - 0002 - 4955 - 3162 -serdarsak@gmail.com. +Sak, Serdar +0000-0002-4955-3162 +Department of Biology, Faculty of Science and Literature, Balıkesir University, Balıkesir, Türkiye. serdarsak @ gmail. com. https: // orcid. org / 0000 - 0002 - 4955 - 3162 +serdarsak@gmail.com. - - -Author + + +Author -Karaytuğ, Süphan -0000-0001-8980-4133 -Department of Biology, Faculty of Science, Mersin University, Mersin, Türkiye. suphankaraytug @ gmail. com. https: // orcid. org / 0000 - 0001 - 8980 - 4133 -suphankaraytug@gmail.com. +Karaytuğ, Süphan +0000-0001-8980-4133 +Department of Biology, Faculty of Science, Mersin University, Mersin, Türkiye. suphankaraytug @ gmail. com. https: // orcid. org / 0000 - 0001 - 8980 - 4133 +suphankaraytug@gmail.com. - - -Author + + +Author -Huys, Rony -0000-0003-2411-7003 -Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. Corresponding author. r. huys @ nhm. ac. uk -r.huys@nhm.ac.uk. +Huys, Rony +0000-0003-2411-7003 +Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. Corresponding author. r. huys @ nhm. ac. uk +r.huys@nhm.ac.uk. -text - - -Zootaxa +text + + +Zootaxa - -2024 - -2024-10-21 + +2024 + +2024-10-21 - -5525 + +5525 - -1 + +1 - -1 -66 + +1 +66 - -http://dx.doi.org/10.11646/zootaxa.5525.1.1 + +http://dx.doi.org/10.11646/zootaxa.5525.1.1 -journal article -10.11646/zootaxa.5525.1.1 -1175-5334 -14021479 -7F2F59B2-E0FB-4E17-BAF1-31228DB9428E +journal article +10.11646/zootaxa.5525.1.1 +1175-5334 +14021479 +7F2F59B2-E0FB-4E17-BAF1-31228DB9428E - + @@ -205,7 +205,7 @@ Northern and (2) Playa Brava in Iquique. - + Remarks. In addition to the ”normal form” (here considered as a distinct species, @@ -256,9 +256,6 @@ speculate that they may represent different species, he preferred to consider th but assigned uncertain status due to the lack of additional morphological information. - -Species incertae sedis - \ No newline at end of file diff --git a/data/62/7E/C6/627EC678F74BFF91FF4EF98E79FCF965.xml b/data/62/7E/C6/627EC678F74BFF91FF4EF98E79FCF965.xml new file mode 100644 index 00000000000..ca8352d626d --- /dev/null +++ b/data/62/7E/C6/627EC678F74BFF91FF4EF98E79FCF965.xml @@ -0,0 +1,309 @@ + + + +Review of Neoleptastacus Nicholls, 1945 (Harpacticoida, Arenopontiidae), including an updated key to species and proposal of Phreatipontia gen. nov. + + + +Author + +Sak, Serdar +0000-0002-4955-3162 +Department of Biology, Faculty of Science and Literature, Balıkesir University, Balıkesir, Türkiye. serdarsak @ gmail. com. https: // orcid. org / 0000 - 0002 - 4955 - 3162 +serdarsak@gmail.com. + + + +Author + +Karaytuğ, Süphan +0000-0001-8980-4133 +Department of Biology, Faculty of Science, Mersin University, Mersin, Türkiye. suphankaraytug @ gmail. com. https: // orcid. org / 0000 - 0001 - 8980 - 4133 +suphankaraytug@gmail.com. + + + +Author + +Huys, Rony +0000-0003-2411-7003 +Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. Corresponding author. r. huys @ nhm. ac. uk +r.huys@nhm.ac.uk. + +text + + +Zootaxa + + +2024 + +2024-10-21 + + +5525 + + +1 + + +1 +66 + + + + +http://dx.doi.org/10.11646/zootaxa.5525.1.1 + +journal article +10.11646/zootaxa.5525.1.1 +1175-5334 +14021479 +7F2F59B2-E0FB-4E17-BAF1-31228DB9428E + + + + +Key to species + + +Redefinition of the generic boundaries in the + +Arenopontiidae ( + +Sak +et al. +2008 + +) + +and addition of new species have outdated the keys by +Lang (1965) +, +Bodiou & Colomines (1986) +, +Karanovic (2000) +and +Wells (2007) +. + +Neoleptastacus accraensis + +and + +N. secundus + +(both +species inquirendae +) are included in +Table 2 +but not in the updated key below. Identifications made with this key must be confirmed by reference to the original descriptions in the literature. + +1. Anal somite with paired dorsolateral spinous spurs or processes................................................2. +Anal somite without paired dorsolateral spurs or processes...................................................13. 2. Abdominal somites with integumental pattern of rectangular plates dorsally and ventrally; P2 enp-2 without inner seta.....3. + +Abdominal somites without conspicuous surface sculpturing; P2 enp-2 with inner seta..............................4. 3. P3 enp-2 with two distal elements; P5 of both sexes with four articulating setae/spines................... + +N. ornamentus +. + + + +P3 enp-2 with one distal element; P5 of both sexes with three articulating setae/spines................... + +N. reductaspina +. + +4. P3 enp-2 with two distal elements........................................................................5. + +P3 enp-2 with one distal element.........................................................................7. + +5. P1 exp-3 with three setae/spines.................................................................... + +N. huysi +. + +P1 exp-3 with four setae/spines..........................................................................6. + + +6. P1 endopod clearly longer than exopod; P3 enp-2 about half the length of enp-1; P5 of both sexes with one seta and two short spines between inner spinous process and outer basal seta............................................ + +N. acanthus +. + +P1 endopod as long as exopod; P3 enp-2 only slightly shorter than enp-1; P5 of both sexes with one seta and one spur between inner spinous process and outer basal seta...................................................... + +N. chaufriassei +. + + + +7. Sternal plates of somites bearing P1–P4 with midventral hook-like processes..................... + +N. emendatus + + +sp. nov. + +Sternal plates of somites bearing P1–P4 without such posteriorly directed processes................................8. + +8. Paired spinous processes on anal somite straight and backwardly directed.........................................9. Paired spinous processes on anal somite dorsally recurved....................................................10. + +9. P1 endopod 1.15 times as long as exopod; P5 elongate in both sexes, length/maximum width ratio 3.0 ( +♀ +) and 3.5 ( +♂ +), respectively................................................................................ + +N. longiremis + +. P1 endopod 1.30 times as long as exopod; P5 much shorter in both sexes, length/maximum width ratio 1.7 ( +♀ +) and 2.25 ( +♂ +), respectively............................................................................. + +N. rectus + + +sp. nov. + + + +10. P1 endopod shorter than exopod........................................................ + +N. abbreviatus + + +sp. nov. + +P1 endopod at least as long as exopod....................................................................11. + + +11. P1 endopod as long as exopod; spinous process on caudal ramus relatively short, about 35% of ramus length.................................................................................................. + +N. chilensis + + +sp. nov. + +P1 endopod 1.25–1.30 times as long as exopod; spinous process on caudal ramus longer, about 45–55% of ramus length..12. + + +12. P1 enp-1 1.25 times as long as enp-2; P +5 ♀ +twice as long as maximum width; terminal process of caudal ramus dorsally recurved.................................................................................... + +N. gussoae + +. P1 enp-1 1.65 times as long as enp-2; P +5 ♀ +2.7 times as long as maximum width; terminal process of caudal ramus not dorsally recurved.................................................................................... + +N. indicus + +. + +13. P1 exopod 2-segmented...............................................................................14. P1 exopod 3-segmented...............................................................................16. + +14. P5 inner spine at least partly delimited at base; P +6 ♂ +with one seta.............................................15. P5 inner spine fused at base, forming spinous process; P +6 ♂ +with two elements.................. + +N. supersetosus + + +sp. nov. + + + +15. P2 exp-2 outer spine shorter than exopod; P2 endopod shorter than exp-1, flask-shaped; P4 exp-3 four times as long as maximum width, with inner seta arising from distal fifth of inner margin......................................... + +N. trisetosus + +. P2 exp-2 outer spine distinctly longer than exopod; P2 endopod as long as exp-1, subrectangular and slightly tapering in distal third; P4 exp-3 2.75 times as long as maximum width, with inner seta arising from middle third of inner margin............................................................................................ + +N. panamensis + + +sp. nov. + + +16. P3 endopod 1-segmented, with one distal element..........................................................17. P3 endopod 2-segmented, enp-2 with two elements (outer one fused at base).....................................19. + +17. P1 exp-3 with four elements; P2 endopod 2-segmented; P4 exp-3 without inner seta....................... + +N. australis +. + +P1 exp-3 with three elements; P2 endopod 1-segmented; P4 exp-3 with inner seta.................................18. + + +18. Inner spinous process of P +5 ♀ +distinctly curved outwardly (unknown in +♂ +); terminal process of caudal ramus dorsally recurved........................................................................ + +N. angolensis + + +comb. nov. + +Inner spinous process of P +5 ♀ +/ +♂ +and terminal process of caudal ramus straight........................... + +N. africanus +. + + + +19. P4 exp-3 without inner seta.................................................................... + +N. pacificus +. + +P4 exp-3 with inner seta...............................................................................20. + + +21. Urosome (except anal somite) with distinct surface ornamentation consisting of elongate rectangular plates............................................................................. + +N. clasingi +. + +Urosome without conspicuous surface ornamentation........................................................22. + +22. Caudal ramus with dorsolateral spur near base of seta VII....................................................23. Caudal ramus without dorsolateral spur near base of seta VII..................................................24. + +23. Lappets of abdominal hyaline frills semi-incised obtusidigitate; inner seta of P2–P3 enp-2 longer than endopod; P5 three times as long as wide, with naked spinous process........................................................ + +N. spicatus + +. Lappets of abdominal hyaline frills denticulate; inner seta of P2–P3 enp-2 shorter than endopod; P5 about 2.5 times as long as wide, with pinnate spinous process.......................................................... + +N. spinicaudatus + +. + + +24. P1 endopod distinctly longer than exopod...................................................... + +N. ishikarianus +. + +P1 rami equally long............................................................ + +N. pseudishikarianus + + +sp. nov. + + + + + \ No newline at end of file diff --git a/data/62/7E/C6/627EC678F74BFF92FF4EFF76780AFA12.xml b/data/62/7E/C6/627EC678F74BFF92FF4EFF76780AFA12.xml new file mode 100644 index 00000000000..b1f679537b7 --- /dev/null +++ b/data/62/7E/C6/627EC678F74BFF92FF4EFF76780AFA12.xml @@ -0,0 +1,283 @@ + + + +Review of Neoleptastacus Nicholls, 1945 (Harpacticoida, Arenopontiidae), including an updated key to species and proposal of Phreatipontia gen. nov. + + + +Author + +Sak, Serdar +0000-0002-4955-3162 +Department of Biology, Faculty of Science and Literature, Balıkesir University, Balıkesir, Türkiye. serdarsak @ gmail. com. https: // orcid. org / 0000 - 0002 - 4955 - 3162 +serdarsak@gmail.com. + + + +Author + +Karaytuğ, Süphan +0000-0001-8980-4133 +Department of Biology, Faculty of Science, Mersin University, Mersin, Türkiye. suphankaraytug @ gmail. com. https: // orcid. org / 0000 - 0001 - 8980 - 4133 +suphankaraytug@gmail.com. + + + +Author + +Huys, Rony +0000-0003-2411-7003 +Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. Corresponding author. r. huys @ nhm. ac. uk +r.huys@nhm.ac.uk. + +text + + +Zootaxa + + +2024 + +2024-10-21 + + +5525 + + +1 + + +1 +66 + + + + +http://dx.doi.org/10.11646/zootaxa.5525.1.1 + +journal article +10.11646/zootaxa.5525.1.1 +1175-5334 +14021479 +7F2F59B2-E0FB-4E17-BAF1-31228DB9428E + + + + + + + +Arenopontia breviarticulata +Mielke, 1975 + + + + + + + + +Arenopontia +( +Neoleptastacus +) +breviarticulata +Mielke, 1975 + +: +Bodin (1979) + + + +Pararenopontia breviarticulata +( +Mielke, 1975 +) +Bodiou & Colomines (1986) + + + + +Arenopontia +( +Pararenopontia +) +breviarticulata +Mielke, 1975 + +: +Bodin (1997) + + +Original description. +Mielke (1975) +:110–112; Abb. 74 ( +♂ +only). + + + + + + +Type +locality. + +Germany +, +Schleswig-Holstein +, +Isle of Sylt +, +List +; in front of old “Litoralstation List/Sylt”; medium to coarse sandy beach + +. + + +Body length. +710 μm ( +♂ +). + + + + +Remarks. +The description of + +Arenopontia breviarticulata + +is based on a single male that displays a 2-segmented P1 exopod (exp-2 and -3 fused) and a reduced P5 bearing only one spiniform and two setiform elements. As pointed out by + +Sak +et al. +(2008) + +it remains unconfirmed whether the inner spine of the male P5 (or spinous process when incorporated in the segment) is genuinely absent or was overlooked.The latter is unlikely given that + +A. breviarticulata + +, with a recorded male body size of 710 μm, is by far the largest arenopontiid to be described so far. + +Sak +et al. +(2008) + +offered an alternative explanation by comparing +Mielke’s (1975) +illustration with the P5 observed in the copepodid IV stage of + +N. indicus + +( +Rao 1967 +: fig. 3-22) which shows a similar underdeveloped condition. This may suggest that + +A. breviarticulata + +has a paedomorphic morphology which is further substantiated by its 2-segmented P1 exopod, a segmentation pattern that is displayed in copepodid II–IV of + +N. indicus + +before a third segment is added at the next moult ( +Rao 1967 +). Conversely, + +A. breviarticulata + +has retained the plesiomorphic armature pattern on P2 endopod and P4 exopod. +Mielke’s (1975) +illustration of the P3 is problematic since it shows an inner seta on exp-3 and two very long setae on enp-2. Such features have not been observed in any other arenopontiid, raising the suspicion that +Mielke (1975) +did not observe the real P3 but duplicated P4 instead. The only difference between his illustrations of P3 and P4 is the extreme disparity in length of the outer basal seta. + + +Mielke (1975) +described the anal operculum with two lateral “Zacken” (jags, sharp projections) which can be regarded as the positional homologues of the paired lateral spinous processes on the anal somite in the + +acanthus + +- group of + +Neoleptastacus + +. The very long outer spines on exp-2 of P2 and P4 (and possibly P3) in + +A. breviarticulata + +differentiates it from all members of this species group. + + +Bodin (1979) +listed + +A. breviarticulata + +under the subgenus +A. +( + +Neoleptastacus + +) while +Mielke (1975) +considered it a member of the subgenus +A. +( + +Arenopontia + +). +Bodiou & Colomines (1986) +placed the species in their new genus + +Pararenopontia + +, the validity of which was dismissed by +Martínez Arbizu & Moura (1994) +on the grounds that it was an artificial taxon uniting species with reduced leg segmentation. + +Sak +et al. +(2008) + +favoured a relationship with the + +Mesopontia +- +Onychopontia +- +Neoleptastacus + +lineage based on the armature formula of P1 enp-2, displaying one geniculate seta and one outer distal spine.Although the morphology of the anal somite suggests that + +A. breviarticulata + +is probably nested within the + +acanthus + +-group of the genus + +Neoleptastacus + +, it is here treated as a +species incertae sedis +in the +Arenopontiidae +and not in + +Neoleptastacus + +as proposed by + +Sak +et al. +(2008) + +. + + + + \ No newline at end of file diff --git a/data/62/7E/C6/627EC678F75AFF83FF4EFA1279F9F82D.xml b/data/62/7E/C6/627EC678F75AFF83FF4EFA1279F9F82D.xml new file mode 100644 index 00000000000..b4a1ffdcff2 --- /dev/null +++ b/data/62/7E/C6/627EC678F75AFF83FF4EFA1279F9F82D.xml @@ -0,0 +1,167 @@ + + + +Review of Neoleptastacus Nicholls, 1945 (Harpacticoida, Arenopontiidae), including an updated key to species and proposal of Phreatipontia gen. nov. + + + +Author + +Sak, Serdar +0000-0002-4955-3162 +Department of Biology, Faculty of Science and Literature, Balıkesir University, Balıkesir, Türkiye. serdarsak @ gmail. com. https: // orcid. org / 0000 - 0002 - 4955 - 3162 +serdarsak@gmail.com. + + + +Author + +Karaytuğ, Süphan +0000-0001-8980-4133 +Department of Biology, Faculty of Science, Mersin University, Mersin, Türkiye. suphankaraytug @ gmail. com. https: // orcid. org / 0000 - 0001 - 8980 - 4133 +suphankaraytug@gmail.com. + + + +Author + +Huys, Rony +0000-0003-2411-7003 +Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. Corresponding author. r. huys @ nhm. ac. uk +r.huys@nhm.ac.uk. + +text + + +Zootaxa + + +2024 + +2024-10-21 + + +5525 + + +1 + + +1 +66 + + + + +http://dx.doi.org/10.11646/zootaxa.5525.1.1 + +journal article +10.11646/zootaxa.5525.1.1 +1175-5334 +14021479 +7F2F59B2-E0FB-4E17-BAF1-31228DB9428E + + + + + + +Key to genera of +Arenopontiidae + + + + + + +1. P1 endopod prehensile, enp-1 distinctly longer than exopod; innermost element of P1 exp-3 penicillate.................................................................................................. + +Arenopontia +Kunz, 1937 + +. + +P1 endopod not prehensile, enp-1 not longer than exopod; innermost element of P1 exp-3 geniculate...................2. +2. P5 of both sexes with innermost element forming a distinct spinous process which is incorporated in the segment.........3. +P5 of both sexes with innermost element not modified (occasionally fused at base).................................7. + +3. Distal margin of P1 enp-2 with two geniculate setae........................................ + +Phreatipontia + + +gen. nov. + + + +Distal margin of P1 enp-2 with outer naked spine and inner geniculate seta............ + +Neoleptastacus +Nicholls, 1945 + +.......4. + + +4. Anal somite with paired spinous processes either side of anal operculum............................. + +acanthus + +-group. + +Anal somite without paired spinous processes either side of anal operculum.......................................5. + +5. P1 exp-1 without outer spine; distal segment of P1 exopod with three elements; P2 endopod 1-segmented... + +trisetosus + +-group. + +P1 exp-1 with outer spine; distal segment of P1 exopod with four elements; P2 endopod 2-segmented..................6. + +6. Distal segment of P3 endopod with one apical element............................................ + +australis + +-group. + + +Distal segment of P3 endopod with two apical elements....................................... + +spinicaudatus + +-group. + + +7. P1 enp-2 with one geniculate seta and one spine; P3 endopod not modified in +♂ +.. + +Mesopontia +Sak, Huys & Karaytuğ, 2008 + +. + +P1 enp-2 with two geniculate setae; P3 endopod sexually dimorphic.............................................8. + +8. P2 enp-2 with inner serrate seta; P3 endopod +♂ +two-segmented...................... + +Psammoleptastacus +Pennak, 1942 + +. + + +P2 enp-2 without inner serrate seta; P3 endopod +♂ +one-segmented............ + +Onychopontia +Sak, Huys & Karaytuğ, 2008 + +. + + + + \ No newline at end of file diff --git a/data/62/7E/C6/627EC678F75AFF83FF4EFD65789FFAAE.xml b/data/62/7E/C6/627EC678F75AFF83FF4EFD65789FFAAE.xml new file mode 100644 index 00000000000..bfccdc4d2b8 --- /dev/null +++ b/data/62/7E/C6/627EC678F75AFF83FF4EFD65789FFAAE.xml @@ -0,0 +1,127 @@ + + + +Review of Neoleptastacus Nicholls, 1945 (Harpacticoida, Arenopontiidae), including an updated key to species and proposal of Phreatipontia gen. nov. + + + +Author + +Sak, Serdar +0000-0002-4955-3162 +Department of Biology, Faculty of Science and Literature, Balıkesir University, Balıkesir, Türkiye. serdarsak @ gmail. com. https: // orcid. org / 0000 - 0002 - 4955 - 3162 +serdarsak@gmail.com. + + + +Author + +Karaytuğ, Süphan +0000-0001-8980-4133 +Department of Biology, Faculty of Science, Mersin University, Mersin, Türkiye. suphankaraytug @ gmail. com. https: // orcid. org / 0000 - 0001 - 8980 - 4133 +suphankaraytug@gmail.com. + + + +Author + +Huys, Rony +0000-0003-2411-7003 +Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. Corresponding author. r. huys @ nhm. ac. uk +r.huys@nhm.ac.uk. + +text + + +Zootaxa + + +2024 + +2024-10-21 + + +5525 + + +1 + + +1 +66 + + + + +http://dx.doi.org/10.11646/zootaxa.5525.1.1 + +journal article +10.11646/zootaxa.5525.1.1 +1175-5334 +14021479 +7F2F59B2-E0FB-4E17-BAF1-31228DB9428E + + + + +Concluding remarks + + + +Neoleptastacus + +is by far the most speciose genus in the family +Arenopontiidae +, currently including 24 valid species. Species identification is hampered by the fact that many members are only known from the +type +locality or a single record, have not been adequately described, or are part of cryptic species complexes ( + +e.g. +gussoae + +-subgroup) which are difficult to unravel.There is little doubt that some species have repeatedly been misidentified ( + +e.g. +N. secundus + +and + +N. indicus + +) while others have mistakenly been treated as conspecific, leading to the misconception that they assume amphi-Pacific or amphi-Panamanian distribution patterns. Although the present review has identified a number of close-knit species groups in + +Neoleptastacus + +, the relationships between them are as yet not fully understood. A forthcoming phylogenetic analysis including all arenopontiid taxa will assess whether some or all of these groups may be accorded generic status or merely reflect diversification within the genus. This analysis will necessarily assess the phylogenetic significance of previously underrated characters such as male antennule morphology (position of geniculation), presence/absence of a middorsal process on the anal operculum and spinulation patterns on the caudal ramus. Some characters may turn out to be potential apomorphies for particular species-(sub)groups such as the presence of a medial spur on caudal ramus in the + +gussoae + +-subgroup, the presence of paired processes on the anal somite ( + +acanthus + +-group) or the fusion of the outer distal spine of P3 enp-2 ( + +spinicaudatus + +-group and others?). Others such as the presence of abdominal integumental patterns ( + +ornamentus + +-subgroup and + +N. clasingi + +) or the shortening of the inner subdistal seta of P4 exp-3 ( + +N. chaufriassei + +and + +N. africanus + +) are clearly the result of convergence that further complicate analysis of character state evolution in the genus and the family. + + + + \ No newline at end of file diff --git a/data/62/7E/C6/627EC678F761FFB7FF4EF9427DE5FEA2.xml b/data/62/7E/C6/627EC678F761FFB7FF4EF9427DE5FEA2.xml new file mode 100644 index 00000000000..7a02323c446 --- /dev/null +++ b/data/62/7E/C6/627EC678F761FFB7FF4EF9427DE5FEA2.xml @@ -0,0 +1,116 @@ + + + +Review of Neoleptastacus Nicholls, 1945 (Harpacticoida, Arenopontiidae), including an updated key to species and proposal of Phreatipontia gen. nov. + + + +Author + +Sak, Serdar +0000-0002-4955-3162 +Department of Biology, Faculty of Science and Literature, Balıkesir University, Balıkesir, Türkiye. serdarsak @ gmail. com. https: // orcid. org / 0000 - 0002 - 4955 - 3162 +serdarsak@gmail.com. + + + +Author + +Karaytuğ, Süphan +0000-0001-8980-4133 +Department of Biology, Faculty of Science, Mersin University, Mersin, Türkiye. suphankaraytug @ gmail. com. https: // orcid. org / 0000 - 0001 - 8980 - 4133 +suphankaraytug@gmail.com. + + + +Author + +Huys, Rony +0000-0003-2411-7003 +Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. Corresponding author. r. huys @ nhm. ac. uk +r.huys@nhm.ac.uk. + +text + + +Zootaxa + + +2024 + +2024-10-21 + + +5525 + + +1 + + +1 +66 + + + + +http://dx.doi.org/10.11646/zootaxa.5525.1.1 + +journal article +10.11646/zootaxa.5525.1.1 +1175-5334 +14021479 +7F2F59B2-E0FB-4E17-BAF1-31228DB9428E + + + + +(3) + +australis + +-group + + + + +Diagnosis. +Anal somite without paired dorsolateral processes. Anal operculum weakly developed, without rounded medial extension. P1 exp-1 with outer spine; exp-3 with four setae/spines. P1 enp-2 with outer spine and inner geniculate seta distally. P2 exp-2 with outer spine of normal length (not extending far beyond distal margin of exp- 3). P2 endopod 2-segmented; enp-2 with inner seta and two distal spines. P3 endopod 1-segmented; with one distal spine. P4 enp-2 with normally developed outer seta. + + + + +Species included. + +N. australis +( +Chappuis, 1953 +) + +. + + + + +This species group includes one incompletely described African species which does not fit comfortably in either the + +spinicaudatus + +- or + +trisetosus + +-groups defined herein. The reduced armature on the P3 endopod (one apical element on the distal segment) is not exclusive to + +N. australis + +( +Table 2 +). More information is required before any statement can be made about its relationships, particularly whether it may be nested as an advanced member in the + +spinicaudatus + +-group. + + + + \ No newline at end of file diff --git a/data/62/7E/C6/627EC678F761FFB8FF4EFBB27EC4F97A.xml b/data/62/7E/C6/627EC678F761FFB8FF4EFBB27EC4F97A.xml new file mode 100644 index 00000000000..a589e8f9da2 --- /dev/null +++ b/data/62/7E/C6/627EC678F761FFB8FF4EFBB27EC4F97A.xml @@ -0,0 +1,181 @@ + + + +Review of Neoleptastacus Nicholls, 1945 (Harpacticoida, Arenopontiidae), including an updated key to species and proposal of Phreatipontia gen. nov. + + + +Author + +Sak, Serdar +0000-0002-4955-3162 +Department of Biology, Faculty of Science and Literature, Balıkesir University, Balıkesir, Türkiye. serdarsak @ gmail. com. https: // orcid. org / 0000 - 0002 - 4955 - 3162 +serdarsak@gmail.com. + + + +Author + +Karaytuğ, Süphan +0000-0001-8980-4133 +Department of Biology, Faculty of Science, Mersin University, Mersin, Türkiye. suphankaraytug @ gmail. com. https: // orcid. org / 0000 - 0001 - 8980 - 4133 +suphankaraytug@gmail.com. + + + +Author + +Huys, Rony +0000-0003-2411-7003 +Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. Corresponding author. r. huys @ nhm. ac. uk +r.huys@nhm.ac.uk. + +text + + +Zootaxa + + +2024 + +2024-10-21 + + +5525 + + +1 + + +1 +66 + + + + +http://dx.doi.org/10.11646/zootaxa.5525.1.1 + +journal article +10.11646/zootaxa.5525.1.1 +1175-5334 +14021479 +7F2F59B2-E0FB-4E17-BAF1-31228DB9428E + + + + +(2) + +spinicaudatus + +-group + + + + +Diagnosis. +Anal somite without paired dorsolateral processes. Anal operculum weakly developed, without rounded medial extension (except for + +A. pseudishikarianus + + +sp. nov. + +). P1 exp-1 with outer spine; exp-3 with four setae/ spines. P1 enp-2 with outer spine and inner geniculate seta distally. P2 exp-2 with outer spine of normal length (not extending far beyond distal margin of exp-3). Endopod P2–P3 2-segmented; P2 enp-2 with inner seta and two distal spines; P3 enp-2 with two distal spines (outer one fused to segment). P4 enp-2 with normally developed outer seta. + + + + +Species included. + +N. spinicaudatus +Nicholls, 1945 + +, + +N. ishikarianus +( +Itô, 1968 +) + +, + +N. clasingi +( +Mielke, 1985 +) + +, + +N. pacificus +( +Mielke, 1985 +) + +, + +N. spicatus +( +Mielke, 1985 +) + +and + +N. pseudishikarianus + + +sp. nov. + + + + + +The + +spinicaudatus + +-group accommodates six morphologically very similar species, all of which are currently restricted to the Pacific Ocean. All species display the plesiomorphic armature on P1–P4, except for + +N. pacificus + +which lacks the inner seta on P4 exp-3 ( +Table 2 +). In the absence of any morphology-based apomorphies that could support the common origin of these species, assessment of the potential monophyly of this species group will have to await the arrival of molecular sequence data. + +Sak +et al. +(2008) + +provided a key to the members of the + +spinicaudatus + +- group (except + +N. pseudishikarianus + + +sp. nov. + +). + + +Chappuis & Rouch’s (1961) +description of + +Neoleptastacus accraensis +( +Lang, 1965 +) + +is inadequate and incomplete. Based on a reinterpretation of the armature pattern of the P3 endopod (see below) the species is treated here as a +species inquirenda +in the + +spinicaudatus + +-group. + + + + \ No newline at end of file diff --git a/data/62/7E/C6/627EC678F763FFB8FF4EF94F7F2BFC2A.xml b/data/62/7E/C6/627EC678F763FFB8FF4EF94F7F2BFC2A.xml new file mode 100644 index 00000000000..5fc0e48741f --- /dev/null +++ b/data/62/7E/C6/627EC678F763FFB8FF4EF94F7F2BFC2A.xml @@ -0,0 +1,780 @@ + + + +Review of Neoleptastacus Nicholls, 1945 (Harpacticoida, Arenopontiidae), including an updated key to species and proposal of Phreatipontia gen. nov. + + + +Author + +Sak, Serdar +0000-0002-4955-3162 +Department of Biology, Faculty of Science and Literature, Balıkesir University, Balıkesir, Türkiye. serdarsak @ gmail. com. https: // orcid. org / 0000 - 0002 - 4955 - 3162 +serdarsak@gmail.com. + + + +Author + +Karaytuğ, Süphan +0000-0001-8980-4133 +Department of Biology, Faculty of Science, Mersin University, Mersin, Türkiye. suphankaraytug @ gmail. com. https: // orcid. org / 0000 - 0001 - 8980 - 4133 +suphankaraytug@gmail.com. + + + +Author + +Huys, Rony +0000-0003-2411-7003 +Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. Corresponding author. r. huys @ nhm. ac. uk +r.huys@nhm.ac.uk. + +text + + +Zootaxa + + +2024 + +2024-10-21 + + +5525 + + +1 + + +1 +66 + + + + +http://dx.doi.org/10.11646/zootaxa.5525.1.1 + +journal article +10.11646/zootaxa.5525.1.1 +1175-5334 +14021479 +7F2F59B2-E0FB-4E17-BAF1-31228DB9428E + + + + +(1) + +acanthus + +-group + + + + +Diagnosis. +Anal somite with paired dorsolateral processes. Anal operculum weakly developed, without medial extension. P1 exp-1 with outer spine; exp-3 with four setae/spines (except + +N. huysi + +). P1 enp-2 with outer spine and inner geniculate seta distally. P2 exp-2 with outer spine of normal length (not extending far beyond distal margin of exp-3). Endopod P2–P3 2-segmented. P2 enp-2 with or without inner seta; with two distal spines. P3 enp-2 with 1–2 distal spines. P4 enp-2 with well developed outer seta. + + + + +TABLE 1. +Valid species and +species inquirendae +(indicated by *) of + +Neoleptastacus +Nicholls, 1945 + +and + +Phreatipontia + +gen. nov. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
Current nameOriginal name and combination
+ +Neoleptastacus +Nicholls, 1945 + + +Neoleptastacus spinicaudatus +Nicholls, 1945 + +
+ +Neoleptastacus australis +( +Chappuis, 1953 +) + + +Neoleptastacus acanthus +( +Chappuis, 1954 +) + + +Neoleptastacus longiremis +( +Chappuis, 1955 +) + + +Neoleptastacus secundus +Krishnaswamy, 1957 + +* + + +Arenopontia australis +Chappuis, 1953 + + +Arenopontia acantha +Chappuis, 1954 + + +Arenopontia longiremis +Chappuis, 1955 + +
+ +Neoleptastacus africanus +( +Chappuis & Rouch, 1961 +) + + +Neoleptastacus accraensis +( +Lang, 1965 +) + +* + +Neoleptastacus indicus +( +Rao, 1967 +) + + +Neoleptastacus ishikarianus +( +Itô, 1968 +) + + +Neoleptastacus gussoae +( +Cottarelli, 1973a +) + + +Neoleptastacus trisetosus +( +Mielke, 1982a +) + + +Neoleptastacus clasingi +( +Mielke, 1985 +) + + +Neoleptastacus pacificus +( +Mielke, 1985 +) + + +Neoleptastacus spicatus +( +Mielke, 1985 +) + + + +Arenopontia africana +Chappuis & Rouch, 1961 + + +Arenopontia accraensis +Lang, 1965 + + +Arenopontia indica +Rao, 1967 + + +Arenopontia ishikariana +Itô, 1968 + + +Arenopontia gussoae +Cottarelli, 1973a + + +Arenopontia trisetosa +Mielke, 1982a + + +Arenopontia clasingi +Mielke, 1985 + + +Arenopontia pacifica +Mielke, 1985 + + +Arenopontia spicata +Mielke, 1985 + +
+ +Neoleptastacus angolensis +( +Bodiou & Colomines, 1986 +) + + +comb. nov. + + +Neoleptastacus chaufriassei +( +Bodiou & Colomines, 1986 +) + + +Neoleptastacus ornamentus +( +Mielke, 1987 +) + + +Neoleptastacus reductaspina +( +Mielke, 1987 +) + + +Neoleptastacus huysi +( +Karanovic, 2000 +) + + + +Arenopontia +( +Neoleptastacus +) +africana + +f. + +angolensis +Kunz, 1971 + + +Arenopontia +( +Neoleptastacus +) +chaufriassei +Bodiou & Colomines, 1986 + + +Arenopontia ornamenta +Mielke, 1987 + + +Arenopontia reductaspina +Mielke, 1987 + + +Arenopontia +( +Neoleptastacus +) +huysi +Karanovic, 2000 + +
+ +Neoleptastacus abbreviatus + + +sp. nov. + + +Neoleptastacus chilensis + + +sp. nov. + + + +Arenopontia +? +gussoae +Cottarelli, 1973a +sensu +Mielke (1982b) + +[partim] + +Arenopontia +? +gussoae +Cottarelli, 1973a +sensu +Mielke (1987) + +[partim] +
+ +Neoleptastacus emendatus + + +sp. nov. + + + +Arenopontia +( +Neoleptastacus +) +acantha accraensis +Lang, 1965 +sensu +Kunz (1971) + +
+ +Neoleptastacus panamensis + + +sp. nov. + + +Neoleptastacus pseudishikarianus + + +sp. nov. + + + +Arenopontia trisetosa +Mielke, 1982a +sensu +Mielke (1982b) + + +Arenopontia +? +ishikariana +Itô, 1968 +sensu +Mielke (1987) + +
+ +Neoleptastacus rectus + + +sp. nov. + + +Neoleptastacus supersetosus + + +sp. nov. + + + +Arenopontia +? +gussoae +Cottarelli, 1973a +sensu +Mielke (1982b) + +[partim] +
+ +Phreatipontia + + +gen. nov. + +
+ +Phreatipontia phreatica + +(Cottarelli, Bruno & Venanzetti, + + +Arenopontia +( +Neoleptastacus +) +phreatica +Cottarelli, Bruno & Venanzetti, 1994 + +
+1994 +) +comb. nov. +
+ +Phreatipontia speluncae + +(Cottarelli, Bruno & + + +Arenopontia +( +Neoleptastacus +) +speluncae +Cottarelli, Bruno & + +
+Venanzetti, 1994) +comb. nov. +Venanzetti, 1994
+ + + +Subgroups included. + +acanthus + +-subgroup, + +gussoae + +-subgroup, + +ornamentus + +-subgroup. + + + + +Remarks. +Members of this group can readily be identified by the presence of paired dorsolateral spinous processes on the anal somite. The presence of these backwardly directed extensions, not found in any other species of the +Arenopontiidae +, is an apomorphic character state supporting the monophyletic status of this lineage. + + +Various authors ( +Itô 1978 +; + +Mielke 1982 +b + +, 1987; +Wells & Rao 1987 +) have recognized a + +gussoae + +-subgroup of morphologically extremely similar species characterized by the presence of only one spine on P3 enp-2: + +N. longiremis +( +Chappuis, 1955 +) + +, + +N. indicus +( +Rao, 1967 +) + +, + +N. gussoae +( +Cottarelli, 1973a +) + +and + +N. sakagamii +( +Itô, 1978 +) + +. + +Neoleptastacus accraensis +( +Lang, 1965 +) + +, which is generally believed to display this character (see below for a reinterpretation), has occasionally been cited as part of this species complex ( + +Mielke 1982 +b + +, 1987). However, the absence of paired anal processes excludes this species from the + +acanthus + +-group. The 1-spine condition on the distal endopodal segment of P3 evolved convergently in the common ancestor of the + +trisetosus + +-group as well as in + +N. australis +( +Chappuis, 1953 +) + +and + +N. reductaspina +( +Mielke, 1987 +) + +( +Table 2 +). +Wells & Rao (1987) +, inspired by the extensive variability observed in their material of + +N. indicus + +, proposed to sink + +N. sakagamii + +as a junior subjective synonym of the latter, but argued that that the caudal ramus and possibly P5 are sufficiently different to maintain + +A. gussoae + +as a distinct species. + +Neoleptastacus secundus + +should also be considered a member of the + +acanthus + +- group since +Krishnaswamy (1957: 97) +clearly stated that “...the anal segment bears two spines posteriorly” (hinted at in his illustration of the male habitus). Unfortunately, the P3 endopod was not figured and his statement that it resembles that of P2 (except for the absence of the inner seta on enp-1) has made most authors assume that P3 enp-2 bears two distal elements ( +Lang 1965 +; +Bodiou & Colomines 1986 +; +Karanovic 2000 +). Given its close similarity to + +N. longiremis + +(see below), which bears one element on this segment, this character must await confirmation. The species is here treated as a +species inquirenda +in the + +gussoae + +-subgroup. Reassessment of + +Mielke’s (1982 +b + +, 1987) descriptions of “ + +Arenopontia +? +gussoae + +” resulted in the recognition of three additional members: + +N. abbreviatus + + +sp. nov. + +, + +N. chilensis + + +sp. nov. + +and + +N. rectus + + +sp. nov. + +Finally, + +Neoleptastacus emendatus + + +sp. nov. + +, proposed here (see below) for + +Arenopontia +( +Neoleptastacus +) +acantha accraensis +Lang, 1965 +sensu +Kunz (1971) + +, also belongs to this subgroup which contains species from the northern Atlantic, Indian Ocean, +Japan +and the Pacific seaboard of South America. + + +The + +ornamentus + +-subgroup currently includes two closely related Chilean species, + +N. ornamentus +( +Mielke, 1987 +) + +and + +N. reductaspina +( +Mielke, 1987 +) + +, that display conspicuous surface ornamentation on the abdominal somites in the form of rectangular plates (areas of integumental reinforcement). Similar surface ornamentation has been reported for + +N. clasingi +( +Mielke, 1985 +) + +in the + +spinicaudatus + +-group. Both + +N. ornamentus + +and + +N. reductaspina + +share the unique absence of the inner seta on P2 enp-2 ( +Table 2 +), giving further credence to the monophyly of this subgroup. + + + +Neoleptastacus acanthus +( +Chappuis, 1954 +) + +, + +N. chaufriassei +( +Bodiou & Colomines, 1986 +) + +and + +N. huysi +( +Karanovic, 2000 +) + +exhibit the plesiomorphic 2-spine condition on P3 enp-2 and are united in the + +acanthus + +-subgroup. No synapomorphy has as yet been identified for this subgroup which includes species from the Mediterranean and the sub-antarctic +Crozet Islands +in the southern Indian Ocean. + + + + \ No newline at end of file diff --git a/data/62/7E/C6/627EC678F76EFFB7FF4EFBCB7CB9F95F.xml b/data/62/7E/C6/627EC678F76EFFB7FF4EFBCB7CB9F95F.xml new file mode 100644 index 00000000000..c187c5a42da --- /dev/null +++ b/data/62/7E/C6/627EC678F76EFFB7FF4EFBCB7CB9F95F.xml @@ -0,0 +1,124 @@ + + + +Review of Neoleptastacus Nicholls, 1945 (Harpacticoida, Arenopontiidae), including an updated key to species and proposal of Phreatipontia gen. nov. + + + +Author + +Sak, Serdar +0000-0002-4955-3162 +Department of Biology, Faculty of Science and Literature, Balıkesir University, Balıkesir, Türkiye. serdarsak @ gmail. com. https: // orcid. org / 0000 - 0002 - 4955 - 3162 +serdarsak@gmail.com. + + + +Author + +Karaytuğ, Süphan +0000-0001-8980-4133 +Department of Biology, Faculty of Science, Mersin University, Mersin, Türkiye. suphankaraytug @ gmail. com. https: // orcid. org / 0000 - 0001 - 8980 - 4133 +suphankaraytug@gmail.com. + + + +Author + +Huys, Rony +0000-0003-2411-7003 +Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. Corresponding author. r. huys @ nhm. ac. uk +r.huys@nhm.ac.uk. + +text + + +Zootaxa + + +2024 + +2024-10-21 + + +5525 + + +1 + + +1 +66 + + + + +http://dx.doi.org/10.11646/zootaxa.5525.1.1 + +journal article +10.11646/zootaxa.5525.1.1 +1175-5334 +14021479 +7F2F59B2-E0FB-4E17-BAF1-31228DB9428E + + + + +(5) + +speluncae + +-group + + + + +Diagnosis. +Anal somite without paired dorsolateral processes. Anal operculum weakly developed, without rounded medial extension. P1 exp-1 with outer spine; exp-3 with 4 setae/spines. P1 enp-2 with two geniculate setae distally. P2 exp-2 with outer spine of normal length (not extending far beyond distal margin of exp-3). Endopod P2–P3 2- segmented; P2 enp-2 with inner seta but only one distal spine. P3 enp-2 with one distal spine. P4 enp-2 outer seta relatively short. + + + + +Species included. + +N. phreaticus +( +Cottarelli, Bruno & Venanzetti, 1994 +) + +, + +N. speluncae +( +Cottarelli, Bruno & Venanzetti, 1994 +) + +. + + + + +Members of this group are restricted to the Mediterranean and the Black Sea where they typically inhabit reduced salinity environments that are exposed to freshwater inflow. They also diverge morphologically from all species currently included in + +Neoleptastacus + +by the armature of the P1 endopod, displaying two geniculate setae on the distal segment instead of an outer spine and an inner geniculate seta. Another shared character is the reduced armature on P2–P3 enp-2, showing only one apical element (0.110 and 0.010, respectively). The + +speluncae + +-group cannot be accommodated in + +Neoleptastacus + +and is here attributed generic rank (see below— + +Phreatipontia + + +gen. nov. + +). + + + + \ No newline at end of file diff --git a/data/62/7E/C6/627EC678F76EFFB7FF4EFE3B7D0EFBF3.xml b/data/62/7E/C6/627EC678F76EFFB7FF4EFE3B7D0EFBF3.xml new file mode 100644 index 00000000000..504c5e15358 --- /dev/null +++ b/data/62/7E/C6/627EC678F76EFFB7FF4EFE3B7D0EFBF3.xml @@ -0,0 +1,180 @@ + + + +Review of Neoleptastacus Nicholls, 1945 (Harpacticoida, Arenopontiidae), including an updated key to species and proposal of Phreatipontia gen. nov. + + + +Author + +Sak, Serdar +0000-0002-4955-3162 +Department of Biology, Faculty of Science and Literature, Balıkesir University, Balıkesir, Türkiye. serdarsak @ gmail. com. https: // orcid. org / 0000 - 0002 - 4955 - 3162 +serdarsak@gmail.com. + + + +Author + +Karaytuğ, Süphan +0000-0001-8980-4133 +Department of Biology, Faculty of Science, Mersin University, Mersin, Türkiye. suphankaraytug @ gmail. com. https: // orcid. org / 0000 - 0001 - 8980 - 4133 +suphankaraytug@gmail.com. + + + +Author + +Huys, Rony +0000-0003-2411-7003 +Department of Life Sciences, Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. Corresponding author. r. huys @ nhm. ac. uk +r.huys@nhm.ac.uk. + +text + + +Zootaxa + + +2024 + +2024-10-21 + + +5525 + + +1 + + +1 +66 + + + + +http://dx.doi.org/10.11646/zootaxa.5525.1.1 + +journal article +10.11646/zootaxa.5525.1.1 +1175-5334 +14021479 +7F2F59B2-E0FB-4E17-BAF1-31228DB9428E + + + + +(4) + +trisetosus + +-group + + + + +Diagnosis. +Anal somite without paired dorsolateral processes. Anal operculum well developed, often produced into a rounded medial extension (not illustrated for + +N. trisetosa + +). P1 exp-1 without outer spine (exopod sometimes 2-segmented); exp-3 with three setae/spines. P1 enp-2 with outer spine and inner geniculate seta distally. P2 exp-2 with very long outer setiform element (extending far beyond distal margin of exp-3). Endopod P2–P3 1-segmented. P2 endopod with inner seta and one distal spine. P3 endopod with one distal spine. P4 enp-2 outer seta reduced. + + + + +Species included. + +N. africanus +( +Chappuis & Rouch, 1961 +) + +, + +N. trisetosus +( +Mielke, 1982a +) + +, + +N. angolensis +( +Bodiou & Colomines, 1986 +) + + +comb. nov. + +, + +N. panamensis + + +sp. nov. + +, + +N. supersetosus + + +sp. nov. + + + + + +The common origin of the species included in this group is strongly supported by five apomorphic character states: (1) absence of outer spine on P1 exp-1, (2) presence of only three elements on the distal segment of P1 exopod, (3) 1-segmented P2–P3 endopods with reduced armature (110 and 010, respectively), (4) P2 exp-2 with very long outer setiform element, extending far beyond distal margin of exp-3, and (5) anal operculum well developed, often produced into a rounded median extension (unconfirmed for + +N. trisetosus + +and + +N. panamensis + + +sp. nov. + +). + +Neoleptastacus africanus + +and + +N. angolensis + + +comb. nov. + +share the plesiomorphic 3-segmented P1 exopod, a distinctly elongate P4 exopod and possibly reduced armature on the P5. + +Neoleptastacus trisetosus + +, + +N. panamensis + + +sp. nov. + +and + +N. supersetosus + + +sp. nov. + +exhibit the apomorphic 2-segmented condition of the P1 exopod. The group has a wide distribution with records from the +Galápagos +archipelago ( +Ecuador +), +Panama +, +Kuwait +and the Atlantic seaboard of Africa. + + + + \ No newline at end of file diff --git a/data/87/00/B5/8700B5F4C3D35D11A71AB708ED412EBD.xml b/data/87/00/B5/8700B5F4C3D35D11A71AB708ED412EBD.xml new file mode 100644 index 00000000000..c90ac6c0b14 --- /dev/null +++ b/data/87/00/B5/8700B5F4C3D35D11A71AB708ED412EBD.xml @@ -0,0 +1,1702 @@ + + + +Two new species of Thomasomys (Cricetidae: Sigmodontinae) from the western Andes of Ecuador and an updated phylogenetic hypothesis for the genus + + + +Author + +Brito, Jorge +0000-0002-3410-6669 +Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador + + + +Author + +García, Rubí +0000-0002-3565-3429 +Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador + + + +Author + +Castellanos, Francisco X. +0000-0003-0955-8185 +Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador & Department of Biological Sciences, Texas Tech University, Lubbock, Texas, United States + + + +Author + +Gavilanes, Gabriela +https://orcid.org/0009-0004-4257-5811 +Laboratorio de Biología Evolutiva, Colegio de Ciencias Biológicas y Ambientales COCIBA, Universidad San Francisco de Quito USFQ, Quito, Cumbayá, Ecuador + + + +Author + +Curay, Jenny +https://orcid.org/0009-0002-4619-0295 +Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador & Instituto de Diversidad y Evolución Austral (IDEAus — CONICET), Puerto Madryn, Argentina + + + +Author + +Carrión-Olmedo, Julio C. +0000-0002-4536-6052 +Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador + + + +Author + +Reyes-Barriga, Daniela +https://orcid.org/0009-0000-4981-9226 +Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador + + + +Author + +Guayasamin, Juan M. +0000-0003-0098-978X +Laboratorio de Biología Evolutiva, Colegio de Ciencias Biológicas y Ambientales COCIBA, Universidad San Francisco de Quito USFQ, Quito, Cumbayá, Ecuador + + + +Author + +Salazar-Bravo, Jorge +0000-0001-5905-2532 +Department of Biological Sciences, Texas Tech University, Lubbock, Texas, United States + + + +Author + +Pinto, C. Miguel +0000-0002-3640-2357 +Charles Darwin Research Station, Charles Darwin Foundation, Puerto Ayora, Galápagos, Ecuador + +text + + +Vertebrate Zoology + + +2024 + +2024-11-05 + + +74 + + +709 +734 + + + +journal article +304986 +10.3897/vz.74.e128528 +71e58ac0-5a79-4833-8211-ce2ff7d2b43b +D2E77D69-6EBF-4736-8557-2E431A6ACCB9 + + + + + +Thomasomys otavalo + +sp. nov. + + + + + +Holotype +. + + + + + +MECN +6912 + +(field number JBM 2574), an +adult female +collected +19 January, 2022 +, by +Jorge Brito +, +Rubí García +and +Fausto Rodríguez +, preserved as dry skin, skull, postcranial skeleton, and muscle and liver biopsies in 95 % ethanol. + + + + + + +Measurements of the +holotype +(in mm, mass in g). + + + +HB += 120; +TL += 161; +HF += 29, E = 19; W = 36; +CIL += 28.62; +LD += 8.49; +LM += 4.7; + +BM +1 + += 1.53; +LIF += 5.54; +BIF += 2.11; +BPB += 3.58; +BZP += 2.54; +LIF += 5.13; +ZB += 16.63; +DI += 1.54; +BIT += 2.03; +LR += 11.12; +LN += 11.84; + +BR + += 5.94; +OL += 10.02; +BMF += 2, 29; +BCO += 7, 54; +BCB += 14.67; +LMN += 19.1; +DR += 3.3. External and craniodental measurements of additional specimens are listed in Table +4 +. + + + + + +Summary of morphometric measurements of all specimens in mm. Species names are accompanied by the number of analyzed individuals between parentheses. Minimum and maximum values are accompanied by mean and standard deviation between parentheses. Abbreviations of characters are detailed in the text. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
- + +T. igor + +sp. nov. +(4) + + +T. otavalo + +sp. nov. +(14) + + +T. silvestris + +(30) + + +T. ucucha + +(22) +
+HB +102–122 (113.75 ± 8.66)110–125 (115.79 ± 4.68)93–138 (109.17 ± 10.65)91–117 (105.93 ± 6.64)
+LT +133–160 (147.25 ± 14.77)150–180 (166.79 ± 8.66)130–160 (146.93 ± 8.05)127–153 (140.54 ± 5.45)
+ +HF + +27–29 (28.25 ± 0.96)27–31 (29.43 ± 1.02)24–30 (27.33 ± 1.63)18–30 (26.31 ± 2.39)
+E +18–20 (19.25 ± 0.96)18–20 (18.86 ± 0.53)16–21 (18.98 ± 1.34)15–28 (19.07 ± 2.38)
+ +CIL + +25.94–27.48 (27.06 ± 0.75)28.05–30.29 (28.86 ± 0.83)25.21–27.66 (26.5 ± 0.55)27.25–29.34 (28.45 ± 0.51)
+ +LD + +7.53–8.54 (8.2 ± 0.47)8.13–9.98 (9.2 ± 0.7)6.65–8.32 (7.71 ± 0.39)8.4–9.94 (9.1 ± 0.34)
+ +LM + +4.74–4.83 (4.8 ± 0.04)4.02–5.44 (4.5 ± 0.4)4.41–5.05 (4.73 ± 0.14)4.29–5.5 (4.57 ± 0.24)
+ +BM 1 + +1.55–1.63 (1.6 ± 0.03)1.41–1.63 (1.52 ± 0.08)1.47–1.68 (1.58 ± 0.06)1.43–1.7 (1.51 ± 0.06)
+ +LIF + +5.53–6.28 (5.86 ± 0.31)5.11–7.09 (5.89 ± 0.77)4.8–5.71 (5.19 ± 0.25)4.97–5.65 (5.25 ± 0.2)
+ +BIF + +1.75–2.21 (1.98 ± 0.22)1.85–2.36 (2.17 ± 0.14)1.81–2.48 (2.12 ± 0.14)1.89–2.32 (2.15 ± 0.12)
+ +BPB + +3.06–3.53 (3.22 ± 0.22)3.45–4.56 (3.96 ± 0.3)2.16–3.78 (3.25 ± 0.31)3.48–4.22 (3.76 ± 0.18)
+ +BZP + +2.14–2.27 (2.17 ± 0.06)2.28–2.93 (2.53 ± 0.16)1.69–2.28 (2.08 ± 0.13)2.36–2.73 (2.56 ± 0.11)
+ +LIB + +5.15–5.21 (5.18 ± 0.03)4.1–5.51 (4.75 ± 0.52)4.8–5.56 (5.09 ± 0.18)4.65–5.17 (4.9 ± 0.12)
+ +ZB + +14.62–15.27 (15.05 ± 0.29)14.55–17.73 (16.19 ± 1.22)14–15.84 (15.12 ± 0.46)16–17.46 (16.69 ± 0.33)
+ +DI + +1.35–1.67 (1.48 ± 0.14)1.43–1.68 (1.53 ± 0.07)– 3.42–2.1 (1.28 ± 0.9)1.51–159 (7.93 ± 31.47)
+ +BIT + +1.62–1.83 (1.77 ± 0.1)1.75–2.14 (1.98 ± 0.11)1.56–1.91 (1.75 ± 0.09)2.08–2.6 (2.24 ± 0.12)
+ +LR + +8.62–10.57 (9.58 ± 0.8)9.71–11.17 (10.4 ± 0.45)9.11–11.89 (10.01 ± 0.73)9.19–10.75 (9.87 ± 0.4)
+ +LN + +10.08–11.61 (11.13 ± 0.71)10.08–11.96 (11.14 ± 0.64)9.55–11.82 (10.68 ± 0.5)10.15–11.57 (10.89 ± 0.45)
+ +BR + +4.96–5.4 (5.19 ± 0.18)5.4–7.07 (6.12 ± 0.7)4.76–5.69 (5.19 ± 0.23)5.39–6.07 (5.77 ± 0.17)
+ +OL + +7.6–9.3 (8.1 ± 0.8)9.27–10.35 (9.96 ± 0.26)7.27–9.65 (8.37 ± 0.77)9.57–10.43 (10.1 ± 0.23)
+ +BCO + +6.83–7.14 (7.04 ± 0.15)7–7.61 (7.24 ± 0.21)6.32–7.46 (7 ± 0.25)6.89–7.4 (7.09 ± 0.13)
+ +BMF + +1.74–2.52 (2.2 ± 0.35)2.01–2.42 (2.24 ± 0.11)1.85–2.68 (2.26 ± 0.18)1.95–2.7 (2.29 ± 0.16)
+ +BCB + +13.31–13.46 (13.39 ± 0.07)10–15.95 (14.23 ± 1.39)10.27–14.31 (13.23 ± 0.8)13.44–14.55 (14.13 ± 0.25)
+ +LMN + +14.35–15.07 (14.84 ± 0.33)14.95–16.64 (15.56 ± 0.42)13.74–16.88 (15.08 ± 0.76)14.91–16.6 (15.98 ± 0.4)
+ +DR + +3.05–3.25 (3.14 ± 0.09)2.89–4.72 (3.47 ± 0.59)2.92–3.6 (3.16 ± 0.13)3.4–3.77 (3.62 ± 0.09)
+ + + + +Type locality. + + +Ecuador +, +Provincia de Imbabura +, +Área de Protección Hídrica Otavalo Mojanda +( + +0.15456 +, +– 78.27536 + +, WGS 84 coordinates taken by GPS at the site of collection, elevation + +3,690 m + +). + + + + + +Paratypes +. + + + + +MECN + +6909, + +MECN + +6911, adult males, all preserved as dry skins and cleaned skulls, collected in +Provincia de Imbabura +, Área de Protección Hídrica Otavalo Mojanda ( +0.15456, – 78.27536 +, +3,680 m +) by J. Brito, M. Yánez and C. Paucar on 19 January, 2022. + +MECN + +6916, adult female, preserved as dry skins and cleaned skull, collected in Área de Protección Hídrica Otavalo Mojanda ( + +0.15824 ° S +, +– 78.28259 + +, + +3,550 +m + +) by J. Brito on 21 January, 2022. + +MECN + +6920, 6921, adult males, all preserved as dry skins and cleaned skulls, collected in Área de Protección Hídrica Otavalo Mojanda ( +0.15456, – 78.27536 +, +3,685 m +) by J. Brito, +R +. García and F. Rodríguez on 21 January, 2022. + +MECN + +5603, juvenile female, + +MECN + +5604, juvenile male, all preserved as dry skins and cleaned skulls, collected in Área Protegida Privada Neblina Norte ( + +0.337535 ° S +, +– 78.432619 + +, + +2,990 +m + +) by J. Brito, J. Curay, E. Beltrán and M. Esparza on 23 March, 2017. + +MECN + +5698, + +MECN + +5700, adult males, + +MECN + +5701, adult female, all preserved as cleaned skulls and the rest of the body in ethanol, collected in Tabla Chupa ( +0.336465, – 78.413066 +, +3,055 m +) by J. Brito, J. Curay, E. Beltrán and M. Esparza on 24 March, 2017. + +MECN + +7869, adult female, + +MECN + +7870, adult male, preserved as dry skin and cleaned skull, collected in Cayapas Chupa ( +0.54117, – 78.48343 +, +3,290 m +) by J. Brito, A. Yacelga on +28 September 2023 +. + +MECN + +7873, adult female, + +MECN + +7874, adult male, preserved as dry skin and cleaned skull; + +MECN + +7877, adult female, + +MECN + +7881, 7882, adult males, all preserved as cleaned skulls and alcoholic bodies, all collected in Cayapas Chupa ( +0.53640, – 78.47506 +, +3,440 m +) by J. Brito, A. Yacelga, L. Rodríguez on +03 October 2023 +. + +MECN + +7871, adult female, preserved as cleaned skulls and alcoholic bodies, collected in +Provincia de Esmeraldas +, Cordillera de Cayapas, Loma Negra ( +0.54574, – 78.49796 +, +3,230 m +) by J. Brito, L. Rodríguez on 01–02 October, 2023. + + + + +Etymology. + + +The specific epithet “ Otavalo ” honors the Otavalo culture, here treated as a noun in apposition. The Otavalo people are recognized for their music and ability for weaving and comercializing textiles. For decades, the Otavalos have been one of the most recognizable and proud indigenous cultures of South America ( +Meisch 2002 +). + + + + +Diagnosis. + + +A species of + +Thomasomys + +unique due to the following combination of characters: Head-body length +110–125 mm +, very long tail (~ 144 of head-body length), with white apical portion ( +15–35 mm +); narrow interorbital region with rounded supraorbital margins; narrow zygomatic arches; long incisive foramina covering approximately ~ 63 % of the diastema, but not extending posteriorly between molar series; upper first molars aligned with posterior margin of the zygomatic plate; subsquamosal fenestra larger than the postglenoid foramen; upper incisors opisthodont. + + + + + +Morphological description of the +holotype +and variation. + + + +Pelage fine, dense, and soft, about +10 – 12 mm +long over the back and rump (Fig. +3 +); somberly colored (dark drab) with low countershading. Dorsal coloration Dark Drab (Color 45) along flanks, shading to Hair Brown (color 277) mid-dorsally. Ventral pelage Medium Neutral Gray (Color 298) basally, with superficial of Ground Cinnamon Medium Neutral Gray (Color 270); not sharply demarcated from dorsal coloration (Fig. +4 A, C, E +). Mystacial vibrissae long, extending beyond pinnae when laid back alongside head. Ears sparsely covered with short, blackish hairs, not contrasting conspicuously with color of head. Hairs over metapodials and digits of manus and pes dark, but with white ungual tufts, abundant and extending well beyond the edge of the claws (Fig. +5 A, B +). Tail longer than the combined length of head and body ( +LT +about 144 % of + +HB + +), uniformly dark, with white apical portion (between +15–35 mm +, Fig. +6 A +); with short hairs, but end with some larger hairs; it is covered with rectangular scales (13 or 14 rows / cm near the base), with three dark brown hispid hairs emerging from the base of each scale, no longer than 1–1.5 scale rows, ventral hairs are whitish in color. Mammae six in inguinal, abdominal, and postaxial pairs. + + + + + + +Live specimen of + +Thomasomys otavalo + + +sp. nov. + +(Cordillera Cayapas, Imbabura, Ecuador), in its natural habitat ( +MECN +7869 paratype). Please note the color of the apical portion of the tail. Photographed by J. Brito on September 28, 2023. + + + + + + + +Dry skin in dorsal ( +A +, +B +), lateral ( +C +– +E +), and ventral ( +E +, +F +) views of + +Thomasomys otavalo + +, + +sp. nov. + +( +A +, +C +, +E +; +MECN +6912, holotype), and + +T. igor + + +sp. nov. + +( +B +, +D +, +F +; +MECN +702, holotype). Scale bar: 20 mm. + + + + + + + +Selected external and soft anatomical traits of + +Thomasomys + +: +A +, +C +dorsal and ( +B +, +D +) plantar surface of right pes ( +A +, +B +: + +T. otavalo + + +sp. nov + +, +MECN +5604, paratype; +C +, +D + +T. igor + + +sp. nov + +, +MECN +708, paratype); +E +soft palate ( + +T. otavalo + + +sp. nov + +, +MECN +6921, paratype); +F +, +G +gross morphology of the stomach ( + +T. otavalo + + +sp. nov + +, +MECN +5700, paratype). Abbreviations: 1–5, digits; b, bordering fold; co, cornified epithelium; d, duodenum; d 1 – d 3, diastemal rugae; ge, glandular epithelium; i, incisura angularis; i 1–15, interdental rugae. Scale bars: 5 mm. + + + + + + + +Distal portion of the tail of some species of + +Thomasomys + +: +A + +T. otavalo + + +sp. nov. + +( +MECN +5604, paratype); +B + +T. ucucha +( +MECN +2691) + +; +C + +T. igor + + +sp. nov. + +( +MECN +708, paratype); +D + +T. silvestris + +( +MECN +5054). Scale bar: 10 mm. + + + +Cranium medium for the genus ( +28.1–30.2 mm +of + +CIL + +). Long rostrum, somewhat acuminate and narrow with the nasal and premaxillary bones extending beyond the anterior face of the incisors (giving the appearance of an incipient rostral tube); poorly developed gnathic process. Posterior margin of the nasal bone not reaching the plane of the lacrimal bone. Shallow zygomatic notches. Small and rounded lacrimal bones. Narrow interorbital region with smooth outer edges, leaving the alveolar maxillary processes exposed in dorsal view (Fig. +7 A +). Anteriorly narrowed zygomatic arches. Supraorbital region with divergent posterior borders (sensu +Steppan 1995 +). Frontoparietal suture “ +V +” - shaped. Broad and rounded braincase, slightly flattened at the outer edges. Small and concave exoccipital. Dorsal profile (in lateral view) distinctively flattened from nasal tips to midfrontal region; anterior margin of zygomatic plate slightly tilted backward. No further development in the ethmoturbinals is distinguished. Small lambdoidal crest. First maxillary molar aligned with the posterior edge of the zygomatic plate (Fig. +8 A +). Zygomatic arches sturdy with long jugals spanning a major segment in the zygomatic process of the maxillary bone. Alisphenoid strut wide and short. Carotid circulation primitive (Pattern 1, sensu +Voss 1988 +), as indicated by the presence of a large stapedial foramen, prominent squamosal-alisphenoid groove, and sphenofrontal foramen. Subsquamosal fenestra larger than the postglenoid foramen; hamular process of squamosal thin, long, slightly curved, and distally applied on the mastoid capsule; tegmen tympani broadly overlapping posterior suspensory process of squamosal. Upper edge of the mastoid not exceeding the edge of the subsquamosal fenestra (Fig. +9 A +). Bullae small; pars flaccida of tympanic membrane present, large; orbicular apophysis of malleus well developed. Paraoccipital process small. Hill foramen very small; long and narrow incisive foramina (averaging 64 % of diastemal length), not approaching M 1. Premaxillary process wide in the middle and narrow at the posterior end, maxillary septum of incisive foramina slender and long. Palate short and broad (sensu +Hershkovitz 1962 +), with mesopterygoid fossa reaching the hypoflexus of M 3. Posterolateral palatal pit small and inconspicuous. Mesopterygoid fossa broad, straight sided, extending anteriorly between third molars; bony roof complete or perforated by narrow, slit-like sphenopalatine openings flanking the presphenoid / basisphenoid suture. Basisphenoid wide with slightly flat edges. Foramen ovale small. Parapterygoid fossae narrow, approximately triangular, with moderately (excavated) anterior limits. Middle lacerate foramen broad. Lateral expressions of parietals present. Auditory bullae small and uninflated with large and wide Eustachian tubes. Dentary moderately long, with long and narrow coronoid process (extends beyond upper edge of condylar process); postcondylar and mental processes poorly developed; deep sigmoid notch. Semilunar recess symmetrical, with lower edge ends pointed. Capsular projection of the root of the incisor inconspicuous. + + + + + + + +Thomasomys otavalo + + +sp. nov. + +(Área de Protección Hídrica Otavalo Mojanda, Imbabura, Ecuador): Cranium in ( +A +) dorsal, ( +B +) ventral, and ( +C +) lateral views, and mandible in ( +D +) labial view ( +MECN +6912 holotype). Scale bar: 10 mm. + + + + + + + +Morphological comparisons. Comparison of diastemal and palate region in four species of + +Thomasomys + +. +A + +T. otavalo + + +sp. nov. + +( +MECN +6912, holotype); +B + +T. ucucha + +( +MECN +7937); +C + +T. igor + + +sp. nov. + +( +MECN +702, holotype); +D + +T. silvestris + +( +MECN +4928). Arrow in (A) indicates that the anterior border of the M 1 at the same level of the posterior end of the zygomatic plate; the lower dotted line indicates the penetration of the mesopterygoid fossa in relation to the hypoflexus of the M 3; abbreviations: hf, hypoflexid; if, incisive foramina; mpf, mesopterygoid fossa; pal, palate; ppp, posterolateral palatal pit; zp, zygomatic plate. Scale bar: 5 mm. + + + + + + + +Alisphenoid-mastoid region comparisons in lateral view in four species of + +Thomasomys + +: +A + +T. otavalo + + +sp. nov. + +( +MECN +6912, holotype); +B + +T. ucucha + +( +MECN +7937); +C + +T. igor + + +sp. nov. + +( +MECN +702, holotype); +D + +T. silvestris + +( +MECN +4928). Arrow in (B) illustrated to upper edge of the mastoid that exceed the edge of the subsquamosal fenestra; while the oblique dotted line indicates the suture between the squamosal and parietal. Abbreviations: hp, hamular process of squamosal; exo, exoccipital; ip, interparietal; pa, parietal; lpp, lateral parietal portion; mas, mastoid; mf, mastoid fenestra; pgf, postglenoid foramen; sq, squamosal; ssf, subsquamosal fenestra; zps, zygomatic process of the squamosal bone. For ease of comparison are not included to scale + + + +Upper incisors large, broad, and opisthodont (Thomas’s angle of 80 °, +Thomas 1919 +; see Fig. +10 A +), with front enamel Light Chrome Orange (color 76); brachydont and pentalophodont molars (sensu +Hershkovitz 1962 +). Parallel upper molars small, pentalophodont, hypsodont in juveniles, and lacking cingula; coronal surfaces crested when unworn; main cusps slightly opposite and sloping backwards when viewed from side (Fig. +11 A +). M 1 rectangular in outline with procingulum divided by anteromedian flexus into unequal anterolabial and anterolingual conules; thin anteroloph; wide mesoloph. M 2 squared in outline; mesoloph showing the same condition as in M 1. M 3 rounded in outline with conspicuous anteroloph; shallow paraflexus; mesolph, metaflexus and hypoflexus distinctive, when unworn. Lower molars with main cusps alternating and sloping backwards when viewed from side. First lower molar (m 1) with short anteromedian flexid that divides the procingulum into subequal anterolabial and anterolingual conules (Fig. +12 A +); ectolophid thin and short mesolophid. Protoflexid of m 2 slim; thin and short mesolophid; m 3 slightly shorter than m 2. + + + + + + +Comparison of the left anterior portion of the rostrum, viewed from left side, in four species of + +Thomasomys + +: +A + +T. otavalo + + +sp. nov. + +( +MECN +6912, holotype); +B + +T. ucucha + +( +MECN +7937); +C + +T. igor + + +sp. nov. + +( +MECN +702, holotype); +D + +T. silvestris + +( +MECN +4928). Thomas’ angles according to incisive and molar planes are indicated as well as the extension of the molar series. Arrow in (A) point to the anterior border of the nasal bone extending beyond the anterior face of the incisors relative to (B); abbreviations: n, nasal; nc, nasolacrimal capsule; p, premaxillary; sf, supraorbital foramen; zp, zygomatic plate. Scale bar: 5 mm + + + + + + + +Comparison of the upper right molar series in occlusal view among some species of + +Thomasomys + +: +A + +T. otavalo + + +sp. nov. + +( +MECN +6912, holotype); +B + +T. ucucha + +( +MECN +7937); +C + +T. igor + + +sp. nov. + +( +MECN +702, holotype); +D + +T. silvestris + +( +MECN +4928); abbreviations: af, anteromedian flexus; al, anteroloph; ml, mesoloph; mfl, metaflexus; pf, paraflexus; prf, protoflexus. Scale bar: 1 mm. + + + + + + + +Comparison of the lower right molar series in occlusal view among some species of + +Thomasomys + +: +A + +T. otavalo + + +sp. nov. + +( +MECN +6912, holotype); +B + +T. ucucha + +( +MECN +7937); +C + +T. igor + + +sp. nov. + +( +MECN +702, holotype); +D + +T. silvestris + +( +MECN +4928); abbreviations: ml, mesolophid; prf, protoflexid. Scale bar: 1 mm. + + +Tuberculum of first rib articulating with transverse processes of seventh cervical and first thoracic vertebrae; second thoracic vertebra with differentially elongated neural spine; vertebral column composed of 19 thoracicolumbar, 16 th with moderately developed anapophyses and 17 th with little developed anapophyses, 4 sacrals (fused), and 42–45 caudal vertebrae; usually the second and third caudal vertebrae with small but complete hemal arches; 12 ribs. + +Soft palate with 3 slightly arched diastemal palatal rugae and 5 pairs of interdentals (Fig. +5 E +). The second and third interdental wrinkles are well arched. The interdental wrinkles are short and leave a conspicuous midline “ channel ”. Large gall bladder present in all specimens examined (n = 11). The stomach is of the unilocular and hemiglandular +type +. The cornified epithelium dominates the corpus and is characterized by its spongy surface; the glandular epithelium is widely distributed over the mostly smooth antrum (Fig. +5 F, G +). The boundary between the two epithelia (= bordering fold, sensu +Carleton 1973 +) is manifested by a thick ridge and exceeds the level corresponding to the esophageal opening to the left. While the angular incisura is barely marked, both the esophageal canal and the angular plica are conspicuous. + + + + +Comparisons. + + + +Thomasomys otavalo + + +sp. nov. + +is most closely related to + +T. ucucha + +, and a candidate species from Zuleta (Fig. +1 +: Part A). To construct this section, we used specimens referred to + +Thomasomys ucucha + +(sensu stricto) near Papallacta, +Provincia de Napo +, +type +locality of the species ( +Voss 2003 +). + +Thomasomys otavalo + + +sp. nov. + +is a medium-sized species (Table +5 +), differing from + +T. ucucha + +(features in parentheses) in having shorter dorsal hairs between +10–12 mm +( +13–15 mm +); tail noticeably long ~ 144 % of +HB +(~ 128 %); white tail tip +15–35 mm +(< +5 mm +). Qualitative craniodental differences between the two species are conspicuous: the nasal and premaxillary bones in + +T. otavalo + + +sp. nov. + +are long, extending well in front of the anterior face of the incisors (short, just beyond the anterior surface of the incisors); upper incisors opisthodont (orthodont, slightly procumbent, see Fig. +10 B +); Thomas angle 80 ° (86 °); M 1 goes beyond the posterior edge of the zygomatic plate (not so); M 1 with short paraflexus and indistinct mesoflexus (short and short); mesolophid of m 1 very short (large). + + + + + + +Selected morphological differences with species that could be confused with the new species, compiled from +Voss (2003) +, +Brito et al. (2019) +and our own observations. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+ +T. otavalo + +sp. nov. + +T. ucucha + +T. igor +sp. nov. + + +T. silvestris + +
+Tail very long (LT about 144 % of +HB +) + +Tail long (LT about 128 % of +HB +) + +Tail long (LT about 130 % of +HB +) + +Tail long (LT about 140 % of +HB +) +
White tail tip (15–35 mm)Tail tip usually not white (when present <5 mm)Tail tip usually not white (when present <5 mm)Tail tip usually not white (when present <5 mm)
Dorsal pelage uniformly grayish brownDorsal pelage uniformly dark grayish brownDorsal pelage verona Brown, darker in the midlineDorsal pelage uniformly dark grayish brown
Incipient rostral tube presentRostral tube absentRostral tube absentRostral tube absent
Thomas angle 80 °Thomas angle 86 °Thomas angle 80 °Thomas angle 80 °
Zygomatic arches converge anteriorlyWidely flaring zygomatic archesZygomatic arches converge anteriorlyZygomatic arches converge anteriorly
Upper first molar aligned with posterior edge of the zygomatic plateFirst upper molar does not reach the posterior edge of the zygomatic plateFirst upper molar does not reach the posterior edge of the zygomatic plateUpper first molar reaches beyond the posterior edge of the zygomatic plate
+Incisive foramina long ( +LIF +about 63 % of +LD +), usually not extending posteriorly between molar alveoli + +Incisive foramina vey short ( +LIF +about 57 % of +LD +), not approaching level of molar alveoli + +Incisive foramina long ( +LIF +about 72 % of +LD +), usually not extending posteriorly between molar alveoli + +Incisive foramina long ( +LIF +about 65 % of +LD +), usually not extending posteriorly between molar alveoli +
Upper edge of the mastoid does not exceed the edge of the subsquamosal fenestraThe superior border of the mastoid goes above the subsquamosal fenestra.Upper edge of the mastoid does not exceed the edge of the subsquamosal fenestraUpper edge of the mastoid does not exceed the edge of the subsquamosal fenestra
Capsular process of lower incisor alveolus usually indistinctCapsular process of lower incisor alveolus usually distinctCapsular process of lower incisor alveolus indistinct or absentCapsular process of lower incisor alveolus indistinct or absent
Upper incisors opisthodont, not procumbentUpper incisors orthodont, conspicuously procumbentUpper incisors opisthodont, not procumbentUpper incisors opisthodont, not procumbent
Subsquamosal fenestra larger than the postglenoid foramenSubsquamosal fenestra usually smaller than postglenoid foramenSubsquamosal fenestra larger than the postglenoid foramenSubsquamosal fenestra larger than the postglenoid foramen
M 3 Paraflexo short and metaflexo indistinctParaflexus and metaflexus shortParaflexus and metaflexus length and fusedParaflexus and mesoflexus short
M 3 comparatively largeM 3 comparatively smallM 3 comparatively largeM 3 comparatively small
Procingulum of m 1 dividedProcingulum of m 1 undividedProcingulum of m 1 dividedProcingulum of m 1 undivided or indistinct
+ + +Other + +Thomasomys +species + +that could be confused with + +T. otavalo + + +sp. nov. + +are + +T. silvestris + +and + +T. igor + + +sp. nov. + +However, + +T. otavalo + + +sp. nov. + +can be differentiated from these species by the white tail tip between +15–35 mm +(< +5 mm +in + +T. silvestris + +and + +T. igor + +). Regarding the cranium, + +T. otavalo + + +sp. nov. + +can be distinguished by the incisive foramina (64 % of + +LD + +), while + +T. silvestris + +and + +T. igor + +are longer (65 and 71 % of +LD +, respectively); in + +T. otavalo + +the alisphenoid strut slender while in + +T. silvestris + +and + +T. igor + +they are wide. As for the molars, in + +T. otavalo + + +sp. nov. + +and in + +T. silvestris + +the M 3 paraflexus short and mesoflexus indistinct, while in + +T. igor + +they are long and fused giving the appearance of a horseshoe. Another species with which + +Thomasomys otavalo + + +sp. nov. + +could be confused is + +Thomasomys vulcani + +(sympatric species), however they are easily distinguished because the new species has noticeably long tail ~ 144 % of +HB +(~ 90 %). + + + + +Distribution. + + +Known so far from less than 10 localities, all between the Otavalo-Mojanda Hydric Protected Area (Área de Protección Hídrica Otavalo Mojanda), from the highlands of Intag in +Provincia de Imbabura +, up to the Cordillera de Cayapas at the provincial boundary between +Esmeraldas +and +Imbabura +, at elevations of + +2,290 +–3,685 +m + +. + +Thomasomys otavalo + + +sp. nov. + +is geographically delimited by the basins of the rivers Mira (north), Guayllabamba (south), to the west by the inter-Andean valley and to the east by the tropical rainforest. + + + + +Natural history. + + + +Thomasomys otavalo + + +sp. nov. + +is, thus far, endemic to the temperate and high Andean zoogeographic areas ( +Albuja et al. 2012 +) of the montane forest ( + +Ministerio del Ambiente del +Ecuador +2013 + +), an environment characterized by trees with abundant orchids, ferns, and bromeliads. + +Thomasomys otavalo + + +sp. nov. + +was collected in mature and disturbed forest where the understory is visually dominated by herbaceous families such as +Poaceae +( + +Chusquea +sp. + +and + +Neurolepis +sp. + +), +Araceae +and +Melastomataceae +. Visually the canopy is dominated by encino ( + +Weinmannia +spp. + +), árbol de papel ( + +Polylepis +spp. + +) and guandera ( + +Clusia +spp. + +). The species was collected in sympatry with the shrew-opossum, + +Caenolestes fuliginosus +(Tomes, 1863) + +, the shrew + +Cryptotis equatoris +( +Thomas, 1912 +) + +, and the rodents + +Akodon mollis +Thomas, 1894 + +, + +Nephelomys moerex +(Thomas, 1914) + +, +Neomicroxus latebricola +(Anthony, 1924), + +Microryzomys minutus +(Tomes, 1860) + +, + +T. vulcani +( +Thomas, 1898 +) + +, + +Thomasomys +sp. + +Mojanda ( + +cinereus + +group), and + +Thomasomys +sp. + +Imbabura-Mojanda ( + +aureus + +group). + + + + \ No newline at end of file diff --git a/data/8D/72/53/8D7253C2E25D570D9C60CC83C0F61931.xml b/data/8D/72/53/8D7253C2E25D570D9C60CC83C0F61931.xml new file mode 100644 index 00000000000..ca76963ebc9 --- /dev/null +++ b/data/8D/72/53/8D7253C2E25D570D9C60CC83C0F61931.xml @@ -0,0 +1,687 @@ + + + +Two new species of Thomasomys (Cricetidae: Sigmodontinae) from the western Andes of Ecuador and an updated phylogenetic hypothesis for the genus + + + +Author + +Brito, Jorge +0000-0002-3410-6669 +Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador + + + +Author + +García, Rubí +0000-0002-3565-3429 +Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador + + + +Author + +Castellanos, Francisco X. +0000-0003-0955-8185 +Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador & Department of Biological Sciences, Texas Tech University, Lubbock, Texas, United States + + + +Author + +Gavilanes, Gabriela +https://orcid.org/0009-0004-4257-5811 +Laboratorio de Biología Evolutiva, Colegio de Ciencias Biológicas y Ambientales COCIBA, Universidad San Francisco de Quito USFQ, Quito, Cumbayá, Ecuador + + + +Author + +Curay, Jenny +https://orcid.org/0009-0002-4619-0295 +Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador & Instituto de Diversidad y Evolución Austral (IDEAus — CONICET), Puerto Madryn, Argentina + + + +Author + +Carrión-Olmedo, Julio C. +0000-0002-4536-6052 +Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador + + + +Author + +Reyes-Barriga, Daniela +https://orcid.org/0009-0000-4981-9226 +Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador + + + +Author + +Guayasamin, Juan M. +0000-0003-0098-978X +Laboratorio de Biología Evolutiva, Colegio de Ciencias Biológicas y Ambientales COCIBA, Universidad San Francisco de Quito USFQ, Quito, Cumbayá, Ecuador + + + +Author + +Salazar-Bravo, Jorge +0000-0001-5905-2532 +Department of Biological Sciences, Texas Tech University, Lubbock, Texas, United States + + + +Author + +Pinto, C. Miguel +0000-0002-3640-2357 +Charles Darwin Research Station, Charles Darwin Foundation, Puerto Ayora, Galápagos, Ecuador + +text + + +Vertebrate Zoology + + +2024 + +2024-11-05 + + +74 + + +709 +734 + + + +journal article +304986 +10.3897/vz.74.e128528 +71e58ac0-5a79-4833-8211-ce2ff7d2b43b +D2E77D69-6EBF-4736-8557-2E431A6ACCB9 + + + + + +Thomasomys igor + +sp. nov. + + + + + + +Thomasomys caudivarius + +– +Salazar-Bravo and Yates (2007 +: fig. 2); partim, not + +Thomasomys caudivarius +Anthony, 1923 + +. + + + + + +Thomasomys caudivarius + +– +Steppan and Schenk (2017 +: fig. 2); partim, not + +Thomasomys caudivarius +Anthony, 1923 + + + + + + + +Thomasomys +sp. 1 + +– +Brito et al. (2019) + +. + + + + + + +Holotype +. + + + + + +MECN +702 + +, an +adult female +collected + +15 July 1994 + +, by +Jorge Salazar-Bravo +(field number JSB 716), originally catalogued at Museum of Southwestern Biology + +MSB + +: Mamm: 70717 ( +NK 27680 +), preserved as dry skin, skull and postcranial skeleton. + + + + + + +Measurements of the +holotype +(in mm, mass in g). + + + +HB += 118; +TL += 160; +HF += 27, E = 19; W = 30; +CIL += 27.44; +LD += 8.49; +LM += 4.74; + +BM +1 + += 1.6; +LIF += 5.77; +BIF += 1.75; +BPB += 3.53; +BZP += 2.14; +LIF += 5.19; +ZB += 15.15; +DI += 1.45; +BIT += 1.81; +LR += 9.51; +LN += 11.61; + +BR + += 5.18; +OL += 7.73; +BMF += 2.43; +BCO += 7, 13; +BCB += 13.31; +LMN += 15.07; +DR += 3.25. External and craniodental of additional specimens are listed in Table +4 +. + + + + +Type locality. + + +Ecuador +, +Provincia de Bolívar +, +Río Tatahuazo +, + +2.5 km +NE de + +Cruz de Lizo +( + +– 1.716667 +, +– 78.98333 + +, WGS 84 coordinates taken by GPS at the collection site, elevation + +2,875 m + +). + + + + + +Paratypes +. + + + + +MECN + +700, juvenile male, + +MECN + +701, adult male, preserved as dry skins and cleaned skulls, by Jorge Salazar-Bravo on +15 July 1994 +. + +MECN + +703, male adult, preserved as dry skins and cleaned skulls, by Nelson Monar. + +MEPN + +6203, adult female preserved as dry skin and cleaned skull, by Igor Castro on +17 and 18 July 1994 +, respectively, collected at +2.5 km +NE de Cruz de Lizo the same locality of the +holotype +. + +MSB + +70712, juvenile female, collected at Cruz de Lizo ( +– 1.716667, – 78.95000 +, +3,000 m +) preserved as dry skins and cleaned skulls, by I. Castro on +18 July 1994 +. + + + + +Etymology. + + +This species is named in honor of Igor A. Castro Revelo (1971–2022), Ecuadorian, prominent collector of rodents and curator of the mammal collection at the Museo Ecuatoriano de Ciencias Naturales ( + +MECN + +) during the period 1994–2001. The species epithet is formed from the name “ Igor ” taken as a noun in apposition. + + + + +Diagnosis. + + +Species of + +Thomasomys + +with a unique combination of characters, as follows: Head-body length +102–122 mm +, with long tail (~ 130 % of head-body length); interorbital region narrow with rounded supraorbital margins; zygomatic arches converging anteriorly; long incisive foramina covering approximately ~ 72 % of the diastema, but not extending posteriorly between the molar series; M 1 without reaching the posterior edge of the zygomatic plate; subsquamosal fenestra larger than the postglenoid foramen; upper incisors opisthodont; M 3 with paraflexus and mesoflexus long and fused; M 3 comparatively large; procingulum of m 1 divided. + + + + + +Morphological description of the +holotype +and variation. + + + +Fine, dense, and soft coat, about +11–13 mm +long on the back and rump. Dorsal coloration Prouts Brown (Color 47), along the flanks changing to Drab (color 19). Ventral coat Medium Neutral Gray (Color 298) basally, with superficial Raw Umber (Color 24); not clearly separated from the dorsal coloration (Fig. +4 B, D, F +). Mystacial vibrissae long, extending beyond the pinnae when placed backward along the head. Ears covered with short, blackish hairs not contrasting with the color of the head. Hairs on metapodials and fingers and toes white and dark giving a gray-haired appearance; ungual tufts white, abundant and extending beyond the edge of the claws (Fig. +5 C, D +). Tail longer than the combined length of head and body (about 130 % of +HB +), uniformly dark, but with white tips (< +5 mm +, Fig. +6 C +); with short sparse hairs, giving a naked appearance at least up to the proximal half, while the distal half is somewhat more hairy; tail end with some larger hairs (< +5 mm +); tail covered with rectangular scales (14 or 15 rows / cm near base), with three dark brown hairs emerging from base of each scale, no longer than 1.5–2 scale rows, ventral hairs with whitish tips. Mammae six in inguinal, abdominal, and postaxial pairs. + + +Skull medium for the genus ( +25.9–27.4 mm + +CIL + +). Rostrum long and narrow (Fig. +13 A +), with nasal and premaxillary bones extending beyond anterior face of incisors; gnathic process poorly developed. Posterior margin of nasal bone reaches and exceeds the plane of the lacrimal bone. Shallow zygomatic notches. Enlarged lacrimal bones, triangular in appearance. Narrow interorbital region with smooth external borders, leaving the maxillary alveolar processes and labial part of the molars exposed in dorsal view. Zygomatic arches narrow anteriorly. Supraorbital region with divergent posterior borders (sensu +Steppan 1995 +). Frontoparietal suture “ +V +” shaped. Broad and rounded braincase, slightly flattened at the outer edges. Broad and concave exoccipital. Dorsal profile (in lateral view) distinctly flattened from nasal tips to middle frontal region; anterior margin of zygomatic plate slightly sloping backward. No further development of the ethmoturbinals. Narrow infraorbital foramen. Not distinguishable lambdoidal crest. M 1 without reaching the posterior edge of the zygomatic plate (Fig. +8 C +). Slender zygomatic arches, with long jugals spanning a similar segment in the zygomatic process of the maxillary and squamosal bone. Alisphenoid strut wide and short. Carotid circulation primitive (Pattern 1, sensu +Voss 1988 +), as indicated by large stapedial foramen, prominent squamosal-alisphenoid groove, and sphenofrontal foramen. Postglenoid foramen approximately twice as large as the subsquamosal fenestra; hamular process of squamosal thin, long, slightly curved and applied distally over the mastoid auditory capsule; tegmen tympani broadly overlapping the posterior suspensory process of squamosal. Upper edge of the mastoid does not exceed the edge of the subsquamosal fenestra (Fig. +9 C +). Bullae small; pars flaccida of tympanic membrane present, large; orbicular apophysis of malleus well developed. Paraoccipital process small. Hill foramen very small; long and narrow incisive foramina (averaging 72 % of diastemal length), not approaching first molar alveoli. Narrow premaxillary process, maxillary septum of the incisive foramen very thin and long. Short and broad palate (sensu +Hershkovitz 1962 +), with the mesopterygoid fossa entering between the molars and reaching the protocone of M 3. Posterolateral palatal pit small and inconspicuous. Wide, slightly divergent-sided mesopterygoid fossa, extending anteriorly between third molars; bony roof perforated by wide, slit-like sphenopalatine openings flanking the basisphenoid. Basisphenoid wide with slightly flat edges. Small foramen ovale. Parapterygoid fossae narrow, approximately triangular, with shallow (unexcavated) anterior limits. Middle lacerate foramen narrow. Lateral expressions of the parietals present, small (Fig. +9 C +). Auditory bullae small and uninflated with short and wide Eustachian tubes. Dentary moderately long, with long and narrow coronoid process (extending beyond upper edge of condylar process); postcondylar and mental processes poorly developed; deep sigmoid notch. Semilunar recess symmetrical, with lower edge ends pointed. Capsular projection of the root of the incisor inconspicuous. + + + + + + + +Thomasomys igor + + +sp. nov. + +(Río Tatahuazo, Bolívar, Ecuador): cranium in ( +A +) dorsal, ( +B +) ventral, and ( +C +) lateral views, and mandible in ( +D +) labial view ( +MECN +702, holotype). Scale bar: 1 mm. + + + +Upper incisors small, slender, and opisthodont (Thomas’s angle of 80 °, +Thomas 1919 +; see Fig. +10 C +), with front enamel Light Chrome Orange (color 76); brachydont and pentalophodont molars (sensu +Hershkovitz 1962 +). Upper molars in left and right parallel series, small and pentalodont; hypsodont and cingulate in juveniles; coronal surfaces crested when unworn; main cusps slightly opposite and inclined backward when viewed from the side. M 1 rectangular in outline with procingulum divided by the anteromedian flexus into subequal anterolabial and anterolingual conules; anteroloph large; mesoloph short and / or segmented. M 2 squared in outline; mesoloph showing the same condition as in M 1. M 3 enlarged relative to M 2, rounded in outline with conspicuous anteroloph; paraflexus and metaflexus long and fused (Fig. +11 C +); mesoloph present in juvenile specimens; hypoflexid conspicuous. Lower molars with alternate, posteriorly inclined main cusps viewed from the side. First lower molar (m 1) with anteromedian flexid dividing procingulum into subequal anterolabial and anterolingual conules (Fig. +12 C +); mesolophid short; mesolophid of m 2 short, hypoflexid deep; m 3 slightly shorter than m 2. + +Tuberculum of first rib articulates with transverse processes of seventh cervical and first thoracic vertebrae; second thoracic vertebra with differentially elongated neural spine; vertebral column composed of 19 thoracolumbar, 4 sacral (fused), and 30–39 caudal vertebrae; usually the second and third caudal vertebrae with small but complete hemal arches; 12 ribs. Details of soft anatomy and genitalia unknown. + + + +Comparisons. + + + +Thomasomys igor + + +sp. nov. + +is retrieved as the sister species to + +T. silvestris + +(Fig. +1 +: Part A). To construct this section, we used specimens referred to + +Thomasomys silvestris + +(sensu stricto) from western +Provincia de Cotopaxi +and +Pichincha +( +Anthony 1924 a +, +1924 b +; +Brito et al. 2019 +). + +Thomasomys igor + + +sp. nov. + +is a medium-sized species (Table +4 +), which differs from + +T. silvestris + +(characteristics in parentheses) by having slightly shorter tail ~ 130 % of +HB +(~ 140 %). Craniodentally, qualitative differences between the two species are conspicuous: M 1 without reaching the posterior edge of the zygomatic plate (beyond); M 3 with paraflexus and metaflexus long and fused (short); procingulum of m 1 divided (undivided or indistinct). + + +Other + +Thomasomys +species + +who could be confused with + +T. igor + + +sp. nov. + +are + +T. otavalo + +and + +T. ucucha + +. However, + +T. igor + + +sp. nov. + +can be differentiated from these species by the white tail tip < +5 mm +( +15–35 mm +in + +T. otavalo + +and <5 + +T. ucucha + +). Regarding the skull, + +T. igor + + +sp. nov. + +can be distinguished by the incisive foramina (72 % of +LD +), while + +T. otavalo + +and + +T. ucucha + +are shorter (63 and 57 % of +LD +, respectively). As for the molars, in + +T. igor + + +sp. nov. + +in M 3 the paraflexus and metaflexus long and fused, while in + +T. otavalo + +and + +T. ucucha + +they are short. + +Thomasomys igor + + +sp. nov. + +was previously referred to as + +T. caudivarius + +by Salazar and Yates (2007), +Pacheco (2015) +, and Steppan and Schenck (2017), however, + +T. caudivarius + +is larger in size (see +Brito et al. 2019 +: table 2) and has a disjunct distribution on the southwestern slope of the Andes. Another species with which + +Thomasomys igor + + +sp. nov. + +could be confused is + +T. aureus + +(sympatric species), however they are easily distinguished because the new species is notoriously smaller ~ 113 % of +HB +(~ 336). + + + + +Distribution. + + + +Thomasomys igor + + +sp. nov. + +is known only from one locality, near to Cruz de Lizo, +Provincia de Bolívar +, in the intersections of the Bosque Protector Cashca Totoras, at elevations of + +2,875 +–3,000 +m + +. The new species is geographically delimited by the basins of the rivers Angamarca (north), Chanchán (south), to the east by the inter-Andean valley and to the west by the tropical rainforest (see Fig. +14 +). + + + + + + +A relief map of Ecuador is depicted with hypsometric tints that show the elevation variations across the region. The sampling localities of five + +Thomasomys +species + +are marked on the map. + + + + + +Natural history. + + + +Thomasomys igor + + +sp. nov. + +is found within the temperate and high Andean zoogeographic areas ( +Albuja et al. 2012 +). The ecosystem corresponds to montane forest ( + +Ministerio del Ambiente del +Ecuador +2013 + +), which is characterized by trees with abundant orchids, ferns and bromeliads. The species was collected in sympatry with the shrew-opossum, + +Caenolestes fuliginosus + +, the shrew + +Cryptotis equatoris + +and the rodents + +Akodon mollis + +, + +Nephelomys albigularis +(Tomes, 1860) + +, and + +Thomasomys aureus +(Tomes, 1860) + +. + + + + \ No newline at end of file