Abstract
Primulinajiulianshanensis F.Wen & G.L.Xu, a new species of Gesneriaceae from Jiulianshan National Nature Reserve of Jiangxi Province, China, is described and illustrated here. Molecular evidence showed it was sister to P.wenii Jian Li & L.J.Yan, while the morphological observation found clear differences between them, petiole, both sides of leaf blades, adaxial surface of the calyx lobes, corolla inside toward the bottom, bract margins covered glandular-pubescent hairs in P.jiulianshanensis (vs. no glandular-pubescent hairs in P.wenii); lateral bracts 4–9 × ca. 2 mm, the central one 2–5 × 1–1.5 mm, adaxially glabrous but sparsely pubescent at apex (vs. lateral bracts 14–16 × 2.5–3.0 mm, the central one 10–12 × 1.3–1.6 mm, all adaxially pubescent); calyx lobes 8–11 × ca. 2 mm, each side with several brown serrate teeth at apex (vs. 14–15 × ca. 2.5 mm, margin entire); filaments and staminodes sparsely yellow glandular-puberulent (vs. white, glabrous).
Keywords: Flora of Jiangxi, Jiulianshan National Nature Reserve, new taxon, Primulinawenii , taxonomy
Introduction
The genus Primulina Hance (Hance 1883) in Gesneriaceae is mainly distributed in the mountainous areas of southern and southwestern China to northern Vietnam, especially in Karst landforms (Wei 2018; Xu et al. 2020b). Since it was redefined (Wang et al. 2011; Weber et al. 2011), many new species have been discovered and published. For example, ten new taxa of Primulina from China were reported in 2020 (Du et al. 2021). As of December 2022, about 240 species (including infraspecies, the same below) have been confirmed throughout the world, of which 224 species are distributed in China. So far, this genus is the largest genus of Gesneriaceae in China (POWO 2022).
Jiangxi Province is located in the mid-subtropical region of East China; the species number of Primulina is not very rich. After consulting references, checking herbarium specimens, and excluding the species that have been mistakenly identified, only 13 species of Primulina have been confirmed in Jiangxi Province (Peng et al. 2021). Three of them were discovered and published after 2011, which are P.lepingensis Z.L.Ning & M.Kang (Ning et al. 2014), P.suichuanensis X.L.Yu & J.J.Zhou (Zhou et al. 2016) and P.inflata Li H.Yang & M.Z.Xu (Xu et al. 2020a).
In April 2021, an interesting population of Primulina was found on a cliff of Danxia landform under the evergreen broad-leaved forest in Jiulianshan National Nature Reserve, Longnan City, Jiangxi Province. Morphologically, this species is similar to P.wenii Jian Li & L.J.Yan (Li et al. 2017) in some characteristics. For example, leaf blades are oblong or broadly rounded, corolla purple, and so on.
We collected the plants at the flowering and fruiting stage in the type locality to make specimens, and at the same time carried out botanical fine anatomical photography to observe carefully. We saw that the indumentum characters of petiole, leaf, bract, calyx, corolla tube, filaments, staminodes of this unknown species were obviously different from P.wenii, and the differences of two species’ calyx lobes and bracts characters can help us easily distinguish them. In order to understand the phylogenetic placements of this unknown species in Primulina, ITS and trnL-F sequences of this species were amplified and included for phylogenetic analysis to examine the relationships of the putative new species.
Materials and methods
Morphological observation
All available specimens of Primulina were used and compared (i. e. those stored in the following herbaria ANU, HITBC, IBK, IBSC, KUN, PE), as was the material of Primulina from recent fieldwork by the authors’ team in South and Southwest China. All the morphological characters, such as leaves, inflorescences, flowers and capsules, were observed and measured in the field. The description, measurements, shape, color and other details given in this description are based on living plants and specimens. We examined distinguished morphological characters of the presumed new species and the compared one, P.wenii, under a dissecting microscope. We described this presumed new species using the terminology of Wang et al. (1990, 1998).
Sampling and DNA sequencing
We randomly selected one plant from the population to collect its leaves for a DNA experiment. Fresh leaf materials were preserved in silica gel for quick drying. Total genomic DNA was extracted from dried leaves using modified cetyltrimethylammonium bromide (CTAB) protocol (Doyle and Doyle 1987). ITS and trnL-F were amplified and sequenced following the methods of Möller et al. (2009) and Smissen et al. (2004), respectively. Besides, we downloaded the ITS and trnL-F sequences from GenBank for 188 Primulina species and two Petrocodon species. Species and GenBank accession numbers employed in this study are listed in Table 1.
Table 1.
Species names and GenBank accession numbers of ITS and trnL-F DNA sequences used in this study.
| Species name | Voucher number | ITS | trnL-F |
|---|---|---|---|
| Petrocodonainsliifolius | CWH88 | KF202291 | KF202298 |
| Petrocodonhancei | CIPeng22903 | KY796057 | KY796059 |
| Primulinaalutacea | YD07 | KY394847 | KY393441 |
| Primulinaargentea | YMBC | KY394848 | KY393442 |
| Primulinabaishouensis | GXLG05 | KY394849 | KY393443 |
| Primulinabalansae | BALAN | MK747141 | MK746274 |
| Primulinabeiliuensis | GXBLBC | KY394850 | KY393444 |
| Primulinabeiliuensisvar.fimbribracteata | SGQJ04 | KY394851 | KY393445 |
| Primulinabicolor | SLHLCB | KY394852 | KY393446 |
| Primulinabipinnatifida | GXLG04 | KY394853 | KY393447 |
| Primulinabobaiensis | BBGL01 | KY394854 | KY393448 |
| Primulinabogneriana | WF7 | MK747166 | MK746225 |
| Primulinabrachytricha | DWDMCZ | KF498048 | KY393450 |
| Primulinabrachytrichavar.magnibracteata | KFC4193 | MK369979 | MK369994 |
| Primulinabrunnea | BRUN | MK747142 | MK746275 |
| Primulinabullata | GXJX06 | KF498071 | KY393451 |
| Primulinacangwuensis | GXLG04 | KY394853 | KY393447 |
| Primulinacardaminifolia | GXLB | MK747131 | MK746255 |
| Primulinacarinata | NTBC | KY394858 | KY393452 |
| Primulinacataractarum | N1 | MW900263 | MW960358 |
| Primulinachizhouensis | JXFY01 | KY394860 | KY393454 |
| Primulinacolaniae | WF8 | MK747167 | MK746224 |
| Primulinaconfertiflora | GDYS05 | MK747101 | MK746253 |
| Primulinacordata | HYH010 | KC190200 | KC190207 |
| Primulinacordifolia | GXRA02 | KY394863 | KY393457 |
| Primulinacordistigma | GDYCXZ | MK747118 | MK746251 |
| Primulinacrassirhizoma | CJGL01 | KY394864 | KY393458 |
| Primulinacrassituba | HNSP | MK747147 | MK746230 |
| Primulinacurvituba | GXHJ01 | MK747137 | MK746242 |
| Primulinadanxiaensis | P22865 | JX506886 | JX506778 |
| Primulinadebaoensis | DBGL01 | KY394868 | KY393462 |
| Primulinadepressa | DXS02 | KY394869 | KY393463 |
| Primulinadiffusa | PJGL01 | KY394871 | KY393465 |
| Primulinadongguanica | DGBC | KY394872 | KY393466 |
| Primulinadrakei | YNCP01 | KY394873 | KY393467 |
| Primulinadryas | HKDMS | KY394875 | KY393469 |
| Primulinaduanensis | DABC | KY394877 | KY393471 |
| Primulinaeburnea | P22908 | JX506891 | JX506783 |
| Primulinaeffusa | KFC4167 | MK369976 | MK369991 |
| Primulinafengkaiensis | KFC4130 | MK369975 | MK369990 |
| Primulinafengshanensis | KFC4195 | MK369970 | MK369985 |
| Primulinafimbrisepala | P22863 | JX506894 | JX506786 |
| Primulinafimbrisepalavar.mollis | GXIB | JX506895 | JX506787 |
| Primulinaflavimaculata | KFC3988 | MK369974 | MK369989 |
| Primulinafloribunda | DHGL01 | KY394886 | KY393480 |
| Primulinafordii | LJM1207202 | MG727881 | MG727878 |
| Primulinafordiivar.dolichotricha | DHS01 | MK747125 | MK746247 |
| Primulinagemella | GEME | MK747146 | MK746254 |
| Primulinaglabrescens | GZLBSM | MK747132 | MK746278 |
| Primulinaglandaceistriata | GXLCHW | MK747114 | MK746256 |
| Primulinaglandulosa | GXPLCG | KY394887 | KY393481 |
| Primulinagongchengensis | GCGL01 | KY394889 | KY393483 |
| Primulinagrandibracteata | YNHK | MK747121 | MK746266 |
| Primulinaguigangensis | GXGGBC | KY394892 | KY393486 |
| Primulinaguihaiensis | GXLG036 | KY394893 | KY393487 |
| Primulinaguizhongensis | GXGZBC | KY394894 | KY393488 |
| Primulinahalongensis | HLW01 | KY394895 | KY393489 |
| Primulinahedyotidea | XWB | JX506905 | JX506797 |
| Primulinaheterochroa | GXMES01 | KY394898 | KY393492 |
| Primulinaheterotricha | HNBT01 | KY394899 | KY393493 |
| Primulinahezhouensis | HZXH | MK747143 | MK746258 |
| Primulinahiepii | WF2 | MK747144 | MK746223 |
| Primulinahochiensis | GXIB | JX506903 | JX506795 |
| Primulinahuaijiensis | GDHJ02 | KF498127 | KY393495 |
| Primulinahuangii | WF12 | MK747138 | MK746231 |
| Primulinahunanensis | Xu11697 | KU220602 | KU220608 |
| Primulinajiangyongensis | HNJY01 | KY394902 | KY393496 |
| Primulinajingxiensis | LZXHGL01 | KY394903 | KY393497 |
| Primulinajiuwanshanica | JWS | MK747116 | MK746260 |
| Primulinajuliae | LJM1210011 | MG727889 | MG727873 |
| Primulinalangshanica | LSCZ | KY394907 | KY393501 |
| Primulinalatinervis | XIN1 | KY394908 | KY393502 |
| Primulinalaxiflora | P22927 | JX506910 | JX506802 |
| Primulinalechangensis | GDLC12 | KY394910 | KY393504 |
| Primulinaleeii | LSGL01 | KY394911 | KY393505 |
| Primulinaleiophylla | GXJX07 | KY394912 | KY393506 |
| Primulinalepingensis | JXLP01 | KY394913 | KY394913 |
| Primulinaleprosa | GXMS055 | KY394914 | KY393508 |
| Primulinalianpingensis | CHLT016 | MH343910 | MH344542 |
| Primulinaliboensis | GXJX08 | KY394917 | KY393511 |
| Primulinaliguliformis | GXIB | JX506912 | JX506804 |
| Primulinalijiangensis | GLS01 | KY394919 | KY393513 |
| Primulinalinearicalyx | KFC4141 | MH032854 | MH032841 |
| Primulinalinearifolia | GXNN01 | KY394921 | KY393515 |
| Primulinalingchuanensis | LCXHGL01 | KY394922 | KY393516 |
| Primulinalinglingensis | LLBC | KY394923 | KY393517 |
| Primulinalinglingensisvar.fragrans | XHLLBC2 | MK746285 | MK746285 |
| Primulinaliujiangensis | LJGL01 | KY394924 | KY393518 |
| Primulinalobulata | GDQX04 | KF498054 | KY393519 |
| Primulinalonggangensis | P22948 | JX506916 | JX506808 |
| Primulinalongicalyx | GXGL01 | KY394927 | KY393521 |
| Primulinalongii | XWB | JX506917 | JX506809 |
| Primulinalongzhouensis | P22963 | JX506918 | JX506810 |
| Primulinalunglinensis | GZXY04 | KY394930 | KY393524 |
| Primulinalunglinensisvar.amblyosepala | LCDE | MK747105 | MK746281 |
| Primulinalungzhouensis | GXJX10 | KY394931 | KY393525 |
| Primulinaluochengensis | LCWCGL01 | KY394932 | KY393526 |
| Primulinalutea | 1844 | JX506921 | JX506813 |
| Primulinalutescens | PBLS01 | MK747135 | MK746263 |
| Primulinalutvittata | KFC4149 | MK369978 | MK369993 |
| Primulinaluzhaiensis | HYH019 | KC190197 | KC190204 |
| Primulinamabaensis | SZY02 | KY394937 | KY393531 |
| Primulinamacrodonta | GXIB | JX506923 | JX506815 |
| Primulinamaculata | Xu11916 | KU220604 | KU220609 |
| Primulinamaguanensis | YNMG | MK747127 | MK746267 |
| Primulinamalipoensis | YNMLP01 | MK747123 | MK746240 |
| Primulinamedica | GXPLCM | KY394940 | KY393534 |
| Primulinamelanofilamenta | GXXA | MK747158 | MK746277 |
| Primulinaminor | WXXH1 | MK747160 | MK746290 |
| Primulinaminutimaculata | GXLZ10 | KY394941 | KY393535 |
| Primulinamoi | SGWY03 | KF498115 | KY393536 |
| Primulinamollifolia | GXESWC | KY394943 | KY393537 |
| Primulinamultifida | DLXHGL01 | KY394946 | KY393540 |
| Primulinanandanensis | GXJX02 | KY393541 | KY393541 |
| Primulinanapoensis | GXIB | JX506930 | JX506821 |
| Primulinaningmingensis | NMGL01 | KY394949 | KY393543 |
| Primulinaobtusidentata | GZJK01 | KF498096 | KY393544 |
| Primulinaophiopogoides | GXFS01 | KF498062 | KY393545 |
| Primulinaorthandra | ZRBC2 | MK747128 | MK746286 |
| Primulinaparvifolia | GGSL01 | KY394952 | KY393546 |
| Primulinapengii | W0397 | KU220603 | KU220610 |
| Primulinapetrocosmeoides | SHDBC | KY394953 | KY393547 |
| Primulinapinnatifida | MS02 | KY394954 | KY393548 |
| Primulinapolycephala | GDLZ06 | KY394955 | KY393549 |
| Primulinaporphyrea | DNGL01 | KU173793 | KU173799 |
| Primulinapseudoeburnea | KY394958 | KY394958 | KY393552 |
| Primulinapseudoglandulosa | GXYS06 | KF498138 | KY393482 |
| Primulinapseudoheterotricha | XWB | JX506933 | JX506824 |
| Primulinapseudolinearifolia | JXY | MK747140 | MK746280 |
| Primulinapseudomollifolia | JMMXH1 | MK747134 | MK746244 |
| Primulinapseudoroseoalba | JFHGL01 | KY394959 | KY393553 |
| Primulinapteropoda | HNCJ01 | KY394960 | KY393554 |
| Primulinapungentisepala | JEGL01 | KY394962 | KY393556 |
| Primulinapurpurea | ZHGL01 | KY394964 | KY393558 |
| Primulinaqingyuanensis | GDQX01 | KY394965 | KY394965 |
| Primulinarenifolia | GXDA02 | KY394966 | KY393560 |
| Primulinarepanda | GXBM03 | KY394968 | KY393562 |
| Primulinaronganensis | GXRA01 | KF498135 | KY393564 |
| Primulinarongshuiensis | GXRS01 | KF498088 | KY393565 |
| Primulinaroseoalba | LDGL01 | KY394972 | KY393566 |
| Primulinarosulata | GXPL05 | KU528874 | KU528884 |
| Primulinarotundifolia | OO3 | KY394975 | KY393569 |
| Primulinarubribracteata | JH01R | KU173791 | KU173797 |
| Primulinasclerophylla | GXDA01 | KY394979 | KY393573 |
| Primulinasecundiflora | GZQZ | MK747119 | MK746279 |
| Primulinashouchengensis | GXYF02 | KY394980 | KY393574 |
| Primulinasichuanensis | SCBC | MK747162 | MK746264 |
| Primulinadryas | HKDMS | KY394875 | KY393469 |
| Primulinasinovietnamica | Peng21956 | MK369973 | MK369988 |
| Primulinaspinulosa | GXFS02 | KF498063 | KY393576 |
| Primulinasubrhomboidea | GXYS02 | KY395018 | KY393577 |
| Primulinasubulata | GDYA01 | KY395020 | KY393579 |
| Primulinasubulatavar.guilinensis | GXHYXH | KY394967 | KY393561 |
| Primulinasubulatisepala | CQAYH01 | MK747122 | MK746246 |
| Primulinasuichuanensis | GDLC07 | KY395021 | KY393580 |
| Primulinaswinglei | GXRX01 | KY395022 | KY393581 |
| Primulinatabacum | LZ01 | KY395023 | KY393582 |
| Primulinatenuifolia | GXBM01 | KY395024 | KY393583 |
| Primulinatenuituba | GZGY01 | KY395025 | KY393584 |
| Primulinatiandengensis | GXTD03 | KY395027 | KY393586 |
| Primulinatribracteata | GXFS04 | KY395028 | KY393587 |
| Primulinatribracteatavar.zhuana | 1877 | JX506952 | JX506843 |
| Primulinatsoongii | ZSGL01 | KY395029 | KY393588 |
| Primulinavaricolor | GXNP01 | KF498086 | KY393589 |
| Primulinaverecunda | LBJX01 | KY395031 | KY393590 |
| Primulinaversicolor | GDYD01 | MK747155 | MK746252 |
| Primulinavestita | QZXT | MK747156 | MK746282 |
| Primulinavillosissima | QXY01 | KY395032 | KY393591 |
| Primulinawenii | WENI | MK747148 | MK746284 |
| Primulinawentsaii | GXLZ047 | KY395033 | KY393592 |
| Primulinawuae | WSBC | MK747159 | MK746265 |
| Primulinaxinningensis | GGGL01 | KY394891 | KY393485 |
| Primulinaxiziae | ZJHZ01 | KY395038 | KY393597 |
| Primulinayangchunensis | GDYC01 | KY395039 | KY393598 |
| Primulinayangshanensis | GDNX01 | KY395040 | KY393599 |
| Primulinayangshuoensis | GXYS07 | KY395042 | KY393601 |
| Primulinayingdeensis | YD03 | KU528876 | KU528886 |
| Primulinayungfuensis | GXIB | JX506957 | JX506848 |
| Primulinazhoui | WF18 | MK747104 | MK746222 |
| Primulinajiulianshanensis | WF217 | OP243287 | OP243283 |
Phylogenetic analysis
We assembled and aligned the newly obtained sequences and those from GenBank using MAFFT v.7.017 (Katoh et al. 2002) and subsequently corrected and combined the ITS and trnL-F sequences in Geneious 9.1.4 (Kearse et al. 2012). We used the Maximum likelihood (ML) and Bayesian inference (BI) analyses to do the phylogenetic analysis of the ITS and trnL-F matrixes, and the combined ITS + trnL-F sequences data-set. The two best supported tree topologies from maximum likelihood (ML) analyses of ITS and trnL-F were visually compared for topological incongruence. A conflict in tree topologies of each tree was considered significant when incongruent topologies both received bootstrap values ≥ 80% (Fu et al. 2022). The ML analyses were conducted using IQ-TREE 1.6.12 (Nguyen et al. 2015) with the GTR+R6 model and 1000 ultrafast bootstrap replicates. For BI analysis, we employed MrBayes v.3.2.6 (Ronquist et al. 2012) to obtain a maximum clade credibility (MCC) tree. Bayesian inference was performed using one million generations, four runs, four chains, a temperature of 0.001, 25% trees discarded as burn-in, and trees sampled every 1,000 generations (1,000 trees sampled in total) with GTR+I+G model.
Results
Phylogenetic analysis
The ITS matrix had a length of 782 characters, with 449 (57.4%) variable characters and 355 (45.4%) parsimony-informative. In comparison, the trnL-F matrix had a length of 836 characters, with 198 (23.6%) variable characters and 93 (11.1%) were parsimony-informative. The comparison of trees for ITS and trnL-F revealed no significant incongruence topology and both indicated that Primulinajiuyishanensis is closely related to P.wenii (Suppl. materials 1, 2). Because the combined dataset resulted in a better-resolved tree with higher support values, we use the combined dataset to do the further molecular studies. The combined data-set was 1628 characters, with 653 (40.1%) variable characters and 452 (27.8%) parsimony-informative, including the indels in all matrixes. The undescribed species and P.wenii were sister groups [Bayesian posterior probabilities (BIPP) = 1.00, ML ultrafast bootstrap support values (UFBoot) = 100%], and we found 10 and 1 different sites in the ITS (totally 615 bp) and the trnL-F (totally 750 bp) sequences between them, respectively (Table 2). belonging to a strongly supported clade (BIPP = 0.99, UFBoot = 98%) included Primulinacrassituba (W.T.Wang) Mich.Möller & A.Weber, P.effusa F.Wen & B.Pan and P.lobulata (W.T.Wang) Mich.Möller & A.Weber (Wang 1982, 1989; Weber et al. 2011; Pan et al. 2017) (Fig. 1).
Table 2.
Sequence differences of ITS and trnL-F regions between Primulinajiulianshanensis and P.wenii.
| Species & marker | Sequences |
|---|---|
| P.jiulianshanensis ITS | 1~CCCGAGAACATGTTTAAAACACGCTTGCGT~30 |
| P.wenii ITS | 1~CCCGAGAACATGTTTAAGACACGCTTGCGT~30 |
| P.jiulianshanensis ITS | 141~CGAGCGCCTCTCCGTCCTGGCTAAGTTCGC~170 |
| P.wenii ITS | 141~CGAGCGCCTCTCCGTACCGGTGAAGTTCGC~170 |
| P.jiulianshanensis ITS | 396~CGTTTTTTCCACGCTCAAAAGGTGTC–GGGGACGA~430 |
| P.wenii ITS | 396~CGTCTTTTCCACGCTCCAAAGGTGTCGGGGGAAGA~430 |
| P.jiulianshanensis trnL-F | 501~GTTCAAAAGTCCTTTATCTT~520 |
| P.wenii trnL-F | 501~GTTCAAAATTCCTTTATCTT~520 |
Figure 1.
Phylogenetic tree of Primulina generated from maximum likelihood (ML) of combined trnL-F and ITS data-set. Numbers of each branch are support values in the order of UFBoot/BIPP. Stars indicate UFBoot = 100% or BIPP = 1.0. The dash (–) indicates a node with UFBoot or < 50% or BIPP < 0.5.
Taxonomic treatment
. Primulina jiulianshanensis
F.Wen & G.L.Xu sp.nov.
F64C620E-DC3C-5C46-87B4-F1EC4700AAA7
urn:lsid:ipni.org:names:77318694-1
Figure 2.
Primulinajiulianshanensis sp. nov. A habitat B population C plant in blooming D habit E adaxial surface of mature leaf blades and petiole F abaxial surface of mature leaf blade and petiole.
Figure 3.
Primulinajiulianshanensis sp. nov. A cyme B adaxial surfaces of bracts C abaxial surfaces of bracts D adaxial surfaces of calyx lobes E abaxial surfaces of calyx lobes F opened corolla G stamens and anthers H one of lateral staminodes I pistil J immature capsules K transverse section of capsule.
Diagnosis.
This new species differs from P.wenii by the combination of the following characteristics: petiole, both sides of leaf blades, adaxial surface of the calyx lobes, corolla inside toward the bottom, bract margins glandular-pubescent (vs. what above-mentioned eglandular-pubescent in P.wenii); lateral bracts 4–9 × ca. 2 mm, the central one 2–5 × 1–1.5 mm, adaxially glabrous but sparsely pubescent at apex (vs. lateral bracts 14–16 × 2.5–3.0 mm, the central one 10–12 × 1.3–1.6 mm, all adaxially pubescent); calyx lobes 8–11 × ca. 2 mm, each side with several brown serrate teeth at apex (vs. 14–15 × ca. 2.5 mm and entire); filaments and staminodes sparsely glandular-puberulent (vs. glabrous). Detailed morphological comparisons with P.wenii are provided in Table 3.
Table 3.
Comparisons between the characters of Primulinajiulianshanensis and P.wenii.
| Characters | Primulinajiulianshanensis | P.wenii |
|---|---|---|
| Petiole indumentum | densely villous and very sparsely glandular pubescent | densely villous |
| Leaf blades indumentum | adaxially densely white to light red villous and very sparsely glandular pubescent, abaxially densely white villous and pubescent, and very sparsely glandular -pubescent | adaxially densely pubescent and villous, abaxially densely appressed pubescent |
| Bract | lateral ones 4–9 × ca. 2 mm, the middle one 2–5 × 1–1.5 mm, all abaxially white or reddish pubescent, adaxially glabrous but sparsely pubescent at apex, margin ciliate and very sparsely glandular-puberulent | lateral ones 14–16 × 2.5–3.0 mm, the central one 10–12 × 1.3–1.6 mm; outside white pubescent and villous, adaxially white pubescent, margin entire and ciliate |
| Calyx lobes | adaxially sparsely white puberulent and glandular puberulent, margin entire but each side of calyx lobes with 1–3 brown crenate teeth at apex; lobes 8–11 × ca. 2 mm | adaxially sparsely shortly pubescent to nearly glabrous, margin entire, lobes 14–15 × ca. 2.5 mm |
| Filament | yellow from middle to base but white upper half, glandular-puberulent | white, glabrous |
| Staminodes | yellowish, lateral ones very sparsely glandular-puberulent | white, glabrous |
Holotype.
China, Jiangxi Province: Ganzhou City, Longnan County, Jiulianshan National Nature Reserve, growing in shady and moist cliffs in the forest, 24°34'8.9"N, 114°25'49.7"E, altitude ca. 440 m, April 20, 2021, Guo-Liang Xu, JLSXGL-20210420 (Holotype IBK!; Isotype: KUN!)
Description.
Herbs perennial, acaulescent, rhizome terete, 1–4 cm long, 0.8–1.5 cm in diam., leaves all basal, 4–8, petiole 10–40 × 4–13 mm, densely villous and very sparsely glandular-pubescent. Leaf blade oblong-elliptic or broadly elliptic, 4–13 × 4–8 cm, thickly chartaceous, more and less fleshly, adaxially pale green to dark green, densely white to light red villous and very sparsely glandular-pubescent, abaxially pale green, densely white villous and pubescent, and very sparsely glandular-pubescent, base apparently asymmetric but cuneate, margin irregularly obtuse-serrate, apex slightly obtuse, lateral veins 4–6 on each side, adaxially inconspicuously sunken, abaxially prominently raised. Inflorescence dichotomous cymes 2–4, axillary, 1–3-branched, 3–9-flowered or more; peduncle and pedicle green, erectly white or light red pubescent; peduncle 5–10 cm long, 2–3 mm in diam.; pedicle 5–15 mm long, 1–2 mm in diam. Bracts 3, lanceolate or spatulate, a pair on either side in same size, opposite, 4–9 mm long, ca. 2 mm wide, the middle one smaller, 2–5 mm long, 1–1.5 mm wide, all three abaxially white or reddish pubescent, adaxially glabrous but sparsely pubescent at apex, margin entire, ciliate and very sparsely glandular-puberulent, apex acute; bracteoles 3, shape and indumentum same as bracts, 2–5 mm long, 1–1.5 mm wide. Calyx 5-parted, lobes lanceolate, 8–11 mm long, ca. 2 mm wide, nearly equal, abaxially densely white or light red villous, adaxially sparsely white puberulent and glandular-puberulent, margin entire but each side of calyx lobes with 1–3 purplish brown crenate at the apex. Corolla pinkish purple to bluish purple, 3.6–4.0 cm long; corolla tube funnelform, 2.6–3 cm long, mouth 1.3–1.6 cm in diam., base ca. 5 mm in diam., outside densely glandular-pubescent, inside from the middle to the base sparsely glandular-puberulent, and the upper part of the corolla tube glabrous; corolla tube abdomen with two obviously longitudinal ridges, the upper part (close to the mouth) of the longitudinal ridge dark bluish purple, and the lower part (close to the bottom) changing into yellowish brown; a dark reddish-brown lump on the upper throat of the corolla tube inside and between upper lip lobes, ovate to spatulate, extending to the middle of the corolla tube, the lump densely glandular-puberulent; a narrow triangular thickened dark reddish-brown stripe extending to the middle of the corolla tube inside at each side of corolla tube and at the junction of the abaxial and adaxial lip; limb distinctly 2-lipped, adaxial lip 2-parted to the middle, lobes broadly ovate to semicircular, apex round, 6–8 mm long, 6–9 mm wide at the bottom; abaxial lip 3-parted to near the base, lobes elliptical to oblong, 8–12 mm long, 7–9 mm wide at the bottom. Stamens 2, adnate to 1.8 cm above the base of corolla tube; filaments linear, yellow from middle to base but white upper half, 8–11 mm long, geniculate near the base, glandular-puberulent; anthers reniform, slightly constricted at the middle, densely villous and fewer glandular-puberulent; staminodes 3, yellowish, lateral ones ca. 6 mm long, adnate to 14 mm above the base of corolla tube, straight, linear, very sparsely glandular-puberulent, apex capitate, the central one ca. 1 mm long, adnate to 5–6 mm above the base of corolla tube, glabrous. Disc annular, ca. 1 mm high, margin undulate, glabrous, white. Pistil pale green, 2.8–3.2 cm long; style linear, 1.9–2.3 cm long, ca. 1 mm in diam., upper part densely glandular-puberulent, lower part densely glandular-puberulent and eglandular-puberulent, ovary oblong, ca. 10 mm long, ca. 2 mm in diam., densely villous and glandular-pubescent, parietal placenta. Stigma acute triangle to narrowly obtrapeziform, 2-lobed, ca. 3 mm long. Capsule linear, 2–4 cm long, parietal placenta, densely villous and glandular-pubescent.
Phenology.
Flowering from April to May, fruiting from June to September.
Etymology.
The specific epithet ‘jiulianshanensis’ is derived from the type locality, Jiulianshan National Natural Reserve, Jiangxi Province, China.
Vernacular name.
九连山报春苣苔 (Chinese name); Jǐu Lían Shān Bào Chūn Jù Tái (Chinese pronunciation).
Distribution and habitat.
We found three small subpopulations in Jiulianshan National Nature Reserve in Jiangxi Province, which are distributed in the shady and wet place on the cliffs under the evergreen broad-leaved forest in the reserve. And the new species is mainly acccompanied by Begoniapalmata D.Don, Utriculariastriatula J.Smith, Selaginellamoellendorffii Hieron., S.involvens (Sw.) Spring, etc.
Conservation status.
At present, only three small subpopulations with total ca. 300 mature individuals of the new species are known in the type locality, Jiulianshan National Natural Reserve, Jiangxi Province, China. The three subpopulations are stable because they are in the reserve. The known AOO and EOO of the new species are about 0.2 km2 and 25 m2, respectively. Thus, if considering its fewer individuals of three subpopulations, it should be temporarily assessed as Near Threated (NT), following the IUCN Red List Categories and Criteria (IUCN Standards and petitions committee 2022).
Notes.
The mainly morphological differences are showed in diagnosis and Table 3. In addition, the insides of the corolla tube are also somewhat different. For example, at the junction of the abaxial and adaxial lip, there are two narrow triangular thickened dark reddish-brown stripes inside the corolla tube in Primulinajiulianshanensis, but there are only two bluish purple spots at the same places in P.wenii; at corolla tube abdomen of P.jiulianshanensis, the upper parts of the longitudinal ridges are dark bluish purple, and the lower parts are yellowish brown, but the longitudinal ridges inside corolla tube of P.wenii are all dark bluish purple; the lump on the upper throat inside corolla tube of P.jiulianshanensis is dark reddish-brown, but P.wenii is dark bluish brown(Li et al. 2017).
The type locality of Primulinawenii is Rixi Township, Fuzhou, Fujian Province, while the type locality of P.jiulianshanensis is Jiulian Mountain, Jiangxi Province. Their type localities are separated by the Wuyi Mountains, and the two places are more than 500 kilometers apart. P.wenii has a narrow distribution range and is only recorded in Rixi Township, Fuzhou, Fujian Province (Li et al. 2017). This area is coastal and the climate zone of this region belongs to the typical subtropical marine monsoon climate. P.wenii grows in the limestone evergreen broad-leaved forest area with stable morphology. P.jiulianshanensis, grows on the rocks under the evergreen broad-leaved forest in Jiulianshan Nature Reserve, which is a typical Danxia landform. The soil forming rock are mainly sandstone and conglomerate in Jiulianshan Nature Reserve, which has a subtropical monsoon humid climate. The morphology of the P.jiulianshanensis in three subpopulations is also relatively stable. Therefore, considering the differences in molecular, morphological, and habitats between P.jiulianshanensis and P.wenii, they should be classified as two different species.
Except for a few species, such as Primulinafimbrisepala (Hand.-Mazz.) Yin Z.Wang, P.eburnea (Hance)Yin Z.Wang, P.tenuituba (W.T.Wang) Yin Z.Wang, P.juliae (Hance) Mich.Möller & A.Weber, most of the species of Primulina are narrowly distributed and endemic. Among the known species worldwide, more than 170 species are endemic to Karst areas in southern to southwestern China and to northern Vietnam (Wei 2018; Xu et al. 2020b). However, the diversity of Primulina in the Danxia landform has not been well understood so far (Yu et al. 2019). For example, the new species, P.suichuanensis, was found in the Danxia landform in Jiangxi Province (Zhou et al. 2016). It should be noted that a new provincial record for Primulinawenii from Jiangxi Province was discovered in Jiulianshan Nature Reserve (Liao et al. 2020). However, we carefully examined the voucher specimen (No. PVHJX-05557, stored in GNNU), and we noted that the voucher was collected from the same subpopulation of P.jiulianshanensis from the same site in Jiulianshan Nature Reserve. Further, we found there are many inconsistencies between the morphological description from the article (Liao et al. 2020) and the corresponding voucher specimens. Thus, this species’ new provincial record of P.wenii in Jiangxi province is a mistaken identification. Lastly, none of the close relatives of Primulina are morphologically similar to this new species found in Jiangxi (Fig. 1). Thus, the new taxon is not easily confused with the others in this province.
Supplementary Material
Acknowledgements
We want to thank Stephen Maciejewski, the Gesneriad Society, and Michael LoFurno, Associate Professor, Temple University, Philadelphia, USA, for their editorial assistance. This study was financially supported by the Basic Research Fund of Guangxi Academy of Sciences (CQZ-C-1901), the Key Sci. & Tech. Research and Development Project of Guangxi (Guike AD20159091 & ZY21195050), the capacity-building project of SBR of CAS (KFJ-BRP-017-68), Basal Research Fund of GXIB (Guizhifa010) and the Fund of Technology Innovation Alliance of Flower Industry (2020hhlm005).
Citation
Xu G-L, Liang L-F, Chen D-Y, Jing Z-F, Zuo X-H, Zuo Z-Y, Wen F (2023) Primulina jiulianshanensis, a new species of Gesneriaceae from Jiangxi Province, China. PhytoKeys 226: 1–16. https://doi.org/10.3897/phytokeys.226.96351
Contributor Information
Zheng-Yu Zuo, Email: zuozhengyu@mail.kib.ac.cn.
Fang Wen, Email: wenfang760608@139.com.
Supplementary materials
The tree of the trnL-F plastid marker for Primulina species
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Author: Zheng-Yu Zuo
Data type
(phylogenetic, genomic, tree)
Explanation note
The trnL-F plastid marker and the ITS nuclear marker separately and discuss any incongruences between the two markers, particularly with respect to the position of the new species.
The tree of the ITS nuclear marker for Primulina species
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Author: Zheng-Yu Zuo
Data type
(phylogenetic, genomic, tree)
Explanation note
The trnL-F plastid marker and the ITS nuclear marker separately and discuss any incongruences between the two markers, particularly with respect to the position of the new species.
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Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Supplementary Materials
The tree of the trnL-F plastid marker for Primulina species
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Author: Zheng-Yu Zuo
Data type
(phylogenetic, genomic, tree)
Explanation note
The trnL-F plastid marker and the ITS nuclear marker separately and discuss any incongruences between the two markers, particularly with respect to the position of the new species.
The tree of the ITS nuclear marker for Primulina species
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Author: Zheng-Yu Zuo
Data type
(phylogenetic, genomic, tree)
Explanation note
The trnL-F plastid marker and the ITS nuclear marker separately and discuss any incongruences between the two markers, particularly with respect to the position of the new species.



