#NEXUS [!This data set was downloaded from TreeBASE, a relational database of phylogenetic knowledge. TreeBASE has been supported by the NSF, Harvard University, Yale University, SDSC and UC Davis. Please do not remove this acknowledgment from the Nexus file. Generated on July 11, 2025; 16:37 GMT TreeBASE (cc) 1994-2008 Study reference: Cohen J.I. 2013. A phylogenetic analysis of morphological and molecular characters of Boraginaceae: Evolutionary relationships, taxonomy, and patterns of character evolution. Cladistics, . TreeBASE Study URI: http://purl.org/phylo/treebase/phylows/study/TB2:S14332] BEGIN TAXA; TITLE Taxa; DIMENSIONS NTAX=224; TAXLABELS Alkanna_orientalis Alkanna_pinardii Alkanna_tinctoria Amsinckia_calycina Amsinckia_spectabilis Amsinckia_tessellata Anchusa_aegyptiaca Anchusa_azurea Anchusa_capellii Anchusa_capensis Anchusa_cespitosa Anchusa_crispa Anchusa_formosa Anchusa_leptophylla Anchusa_officinalis Anchusa_pusilla Anchusa_stylosa Anchusa_thessala Anchusa_undulata Anchusella_variegata Antiphytum_floribundum Antiphytum_hintoniorum Arnebia_benthami Arnebia_guttata Asperugo_procumbens Borago_morisiana Borago_officinalis Borago_pygmaea Borago_trabutii Bothriospermum_tenellum Bourreria_succulenta Brachybotrys_paridiformis Brandella_sp. Brunnera_orientalis Buglossoides_arvensis Buglossoides_incrassata 'Buglossoides purpureo-caeruleum' Buglossoides_tenuiflora Caccinia_strigosa Cerinthe_alpina Cerinthe_major Chionocharis Codon_schenkii Cordia_boissieri Cordia_dentata Cordia_myxa Cordia_sinensis Cryptantha_C308 Cryptantha_bakeri Cryptantha_confertiflora Cryptantha_crassipes Cryptantha_crassisepala Cryptantha_fendleri Cryptantha_flava_C389 Cryptantha_flavoculata Cryptantha_fulvocanescens Cryptantha_paysonii Cryptantha_peruviana Cryptantha_pterocarya Cynoglossopsis_sp. Cynoglossum_LHMS Cynoglossum_amabile Cynoglossum_javanicum Cynoglossum_officinale Cynoglossum_pringlei Cynoglottis_barrelieri Cynoglottis_chetikiana Cystostemon_heliocharis Dasynotus_sp. Echiochilon_callianthum Echiochilon_fruticosum Echiochilon_johnstonii Echiochilon_longiflorum Echiostachys_incanus Echium_aculeatum Echium_angustifolium Echium_bonnetii Echium_candicans Echium_decaisnei Echium_italicum Echium_leucophaeum Echium_plantagineum Echium_russicum_Pontechium_maculatum Echium_vulgare Echium_wildpretii Ehretia_cymosa Elizaldia_calycina Eritrichium_aretioides Eritrichium_caucasicum Eritrichium_chamissonis Eritrichium_nanum Eritrichium_rupestre Eritrichium_sericeum_subsp._sericeum Eritrichium_splendens Eritrichium_villosum Gastrocotyle_macedonica Glandora_diffusa Glandora_oleifolia Glandora_rosmarinifolia Greeneocharis_circumscissa Hackelia_floribunda Hackelia_micrantha Hackelia_virginiana Halacsya_sendtneri Heliotropium_aegyptiacum Heliotropium_longistylum Hormuzakia_aggregata Huynhia_pulchra Hydrophyllum_canadense Lappula_redowskii Lappula_squarrosa Lappula_texana Lasiocaryum_munroi Lepechiniella_sp. Lindelofia_longiflora Lindelofia_macrostyla Lithodora_hispidula Lithodora_zahnii Lithospermum_bejariense Lithospermum_distichum Lithospermum_incisum Lithospermum_leonotis Lithospermum_macromeria Lithospermum_multiflorum Lithospermum_nelsonii Lithospermum_officinale Lobostemon_fruticosus_48 Lobostemon_trigonus Lycopsis_arvensis Macrotomia_densiflora Maharanga_emodi Mairetis_microsperma Melanortocarya_sp. Mertensia_asiatica Mertensia_ciliata Mertensia_longiflora Mertensia_paniculata_C405 Mertensia_sp._C282 Mertensia_virginica Moltkia_angustifolia Moltkia_aurea Moltkia_caerulea Moltkia_petraea Moltkia_suffruticosa Moltkiopsis_ciliata Moritzia_lindenii Myosotidium_hortensia Myosotis_alpestris Myosotis_arvensis Myosotis_australis Myosotis_discolor Myosotis_macrantha Myosotis_macrosperma Myosotis_sylvatica Neatostema_apulum Nonea_lutea Nonea_pulla Nonea_stenosolen Ogastemma_pusillum Omphalodes_aliena Omphalodes_cappadocica Omphalodes_lojkae Omphalodes_nitida Omphalodes_verna Onosma_alborosea Onosma_echioides Onosma_graecum Onosma_stellulata Onosma_taurica Onosma_visianii Oreocarya_cana Paracaryum_intermedium Paracaryum_lithospermifolium Paracaryum_racemosum Paramoltkia_doerfleri Paraskevia_cesatiana Pardoglossum_cheirifolium Pectocarya_anomala Pectocarya_penicillata_ITS Pectocarya_peninsularis Pentaglottis_sempervirens Phacelia_tanacetifolia Phyllocara_aucheri Plagiobothrys_albiflorus Plagiobothrys_kingii Plagiobothrys_myosotoides Podonosma_orientalis Procopiania_cretica Pseudomertensia_echioides Pseudomertensia_moltkioides_subsp._moltkioides Pseudomertensia_primuloides Pseudomertensia_trollii Pulmonaria_angustifolia Pulmonaria_obscura Pulmonaria_officinalis Rochelia_cancellata Rochelia_persica Solenanthus_apenninus Solenanthus_circinatus Solenanthus_stamineus Suchteliana_sp. Symphytum_asperum Symphytum_caucasicum Symphytum_ibericum Symphytum_orientale Symphytum_peregrinum Symphytum_tuberosum Thaumatocaryon_dasyanthum Thaumatocaryon_tetraquetrum Tiquilia_darwinii Tiquilia_paronychioides Tournefortia_laurifolia Trachystemon_orientalis Tricardia_watsonii Trichodesma_africanum Trichodesma_scotti Trigonocaryum_involucratum Trigonotis_formosana Trigonotis_guilielmii Trigonotis_peduncularis Tysonia_africana Ulugbekia_tschimganica Vahlia_capensis Wellstedia_dinteri ; END; BEGIN TREES; TITLE Boraginaceae_Phyl_Combined; LINK TAXA = Taxa; TREE t1 = [&R] ((Vahlia_capensis:0.07596903,((Hydrophyllum_canadense:0.02401043,Phacelia_tanacetifolia:0.02727145,Tricardia_watsonii:0.02670841):0.01130995,((Bourreria_succulenta:0.03395132,(Ehretia_cymosa:0.01850684,(Tiquilia_darwinii:0.0087315,Tiquilia_paronychioides:0.01013558):0.05550205):0.00266287):0.01178221,(Cordia_boissieri:0.02872714,(Cordia_dentata:0.01414513,(Cordia_sinensis:0.01146825,Cordia_myxa:0.00627189):0.00576525):0.02142657):0.02246105,(Heliotropium_aegyptiacum:0.02329746,Tysonia_africana:0.12506316,(Tournefortia_laurifolia:0.00987752,Heliotropium_longistylum:0.01398093):0.00954482):0.02068407):0.00501852):0.00843805):0.007518095,(Codon_schenkii:0.03379439,(Wellstedia_dinteri:0.05563544,(((Ogastemma_pusillum:0.00989218,(Antiphytum_floribundum:0.00350569,Antiphytum_hintoniorum:0.02018535):0.01622452):0.00321494,(Echiochilon_fruticosum:0.03316382,(Echiochilon_callianthum:0.07476672,(Echiochilon_johnstonii:0.00895223,Echiochilon_longiflorum:0.02147423):0.0224477):0.02411568):0.02276825):0.00858984,(((Chionocharis:0.04619512,Lasiocaryum_munroi:0.07402333):0.04882383,(Caccinia_strigosa:0.01447252,(Trichodesma_africanum:0.00991334,Trichodesma_scotti:0.01150233):0.00342171):0.0068977,((Omphalodes_aliena:0.01948541,(Myosotidium_hortensia:0.00777706,(Omphalodes_lojkae:0.0135705,(Omphalodes_nitida:0.00525501,(Omphalodes_cappadocica:0.0013144,Omphalodes_verna:9.66E-4):0.00151971):0.00358831):0.00243838):7.9041E-4):0.01280178,((Asperugo_procumbens:0.02228628,(Mertensia_asiatica:0.00346458,(Mertensia_sp._C282:0.00415942,(Mertensia_ciliata:0.00244144,Mertensia_virginica:0.00254476):6.7485E-4,(Mertensia_longiflora:0.00209056,Mertensia_paniculata_C405:0.00549888):0.00909997):0.00257131):0.00803906):0.00741406,((Suchteliana_sp.:0.02655469,(Rochelia_cancellata:0.00225219,Rochelia_persica:0.00381884):0.01956138,((Hackelia_micrantha:0.00246433,(Hackelia_virginiana:1.0E-8,Hackelia_floribunda:0.00283892):0.00128765):0.0078579,((Lappula_texana:0.00881142,(Lepechiniella_sp.:0.01027974,Lappula_redowskii:0.00549432,Lappula_squarrosa:0.0010706):0.00230037):0.00730245,(Eritrichium_rupestre:0.00863088,((Eritrichium_sericeum_subsp._sericeum:0.00518389,Eritrichium_splendens:0.00330756):0.01024455,(Eritrichium_caucasicum:0.00968902,(Eritrichium_villosum:9.4728E-4,Eritrichium_nanum:0.00308677,(Eritrichium_aretioides:0.00292792,Eritrichium_chamissonis:0.00456849):0.00170475):0.00162984):0.00787009):0.00816451):0.00244783):0.00184977):0.00455758):0.01053586,(((Trigonocaryum_involucratum:0.01545488,(Myosotis_arvensis:1.0E-8,Myosotis_sylvatica:7.6675E-4):0.0099163,(Myosotis_alpestris:0.01579837,(Myosotis_discolor:0.00911118,(Myosotis_macrosperma:0.00969382,(Myosotis_australis:0.0016526,Myosotis_macrantha:1.0E-8):0.00614686):0.00149277):0.00938655):0.00678194):0.00825641,((Pseudomertensia_trollii:0.00672156,Pseudomertensia_moltkioides_subsp._moltkioides:0.00294862,(Pseudomertensia_primuloides:0.01457125,Pseudomertensia_echioides:0.0087546):0.00153045):0.01044112,(Brachybotrys_paridiformis:0.00768997,Trigonotis_formosana:0.0174976,Trigonotis_guilielmii:0.00157167,Trigonotis_peduncularis:0.04740903):0.01362947):0.00685609):0.00657836,((Bothriospermum_tenellum:0.05677627,(Paracaryum_intermedium:0.01338192,(((Lindelofia_longiflora:0.00245429,Lindelofia_macrostyla:0.00489545):5.4131E-4,(Brandella_sp.:0.03191637,Cynoglossum_LHMS:0.00502427,Cynoglossopsis_sp.:0.02332777,(Cynoglossum_amabile:0.01963755,Cynoglossum_javanicum:0.00387888):0.00232484):4.5746E-4):0.004963,((Solenanthus_apenninus:0.01464702,(Solenanthus_circinatus:0.00225493,Solenanthus_stamineus:0.010449):0.00941556):0.00826775,((Cynoglossum_officinale:0.00232296,Pardoglossum_cheirifolium:0.00786508):0.0017006,(Paracaryum_lithospermifolium:0.00293652,Paracaryum_racemosum:0.00195925):6.5434E-4):0.00101127):9.8064E-4):0.00527284):0.00748001):0.00326885,(Cynoglossum_pringlei:0.00224042,(((Cryptantha_pterocarya:0.02171978,(Cryptantha_crassisepala:0.0093996,Cryptantha_peruviana:0.01055682):0.00202137):0.00190258,(Oreocarya_cana:0.00160239,(Cryptantha_C308:9.8213E-4,(Cryptantha_crassipes:0.00151867,Cryptantha_paysonii:0.00176537):0.00243437,(Cryptantha_bakeri:5.4651E-4,Cryptantha_flavoculata:0.00143718):0.00126013):0.00371562):0.00397583):0.00111401,((Dasynotus_sp.:0.01607595,(Cryptantha_fulvocanescens:0.0018616,(Cryptantha_flava_C389:0.00183029,Cryptantha_confertiflora:5.8822E-4):4.1202E-4):0.00350751):0.0044772,((Plagiobothrys_kingii:0.0124913,(Amsinckia_spectabilis:0.0105222,(Amsinckia_tessellata:0.00802352,Amsinckia_calycina:0.01261789):0.00145285):0.00374063):0.00149309,((Pectocarya_anomala:0.00835439,(Pectocarya_peninsularis:0.00316051,Pectocarya_penicillata_ITS:0.01404542):0.0054755):0.01911287,(Greeneocharis_circumscissa:0.02865858,(Cryptantha_fendleri:0.04636233,(Plagiobothrys_albiflorus:0.01071046,Plagiobothrys_myosotoides:0.0182214):0.00872985):0.00384039):0.00423516):0.00276682):0.00390436):7.7978E-4):0.00293087):0.00558788):0.00814686):0.00527292):0.00552922):0.00181026):0.00759263):0.01025447,(((Melanortocarya_sp.:0.01628212,Nonea_pulla:0.00920063,Paraskevia_cesatiana:0.02068018,(Elizaldia_calycina:0.02474776,(Nonea_lutea:0.00484095,Nonea_stenosolen:0.00317232):0.00941908):0.00534867,(Pulmonaria_angustifolia:0.00939103,(Pulmonaria_obscura:0.00104733,Pulmonaria_officinalis:0.00228321):0.00189346):0.01168281):0.01345176,((Trachystemon_orientalis:0.01464097,((Borago_trabutii:0.01639189,(Borago_officinalis:0.00943433,(Borago_morisiana:0.00427119,Borago_pygmaea:0.00291651):0.01275182):0.00746656):0.01267366,(Pentaglottis_sempervirens:0.01226606,(Procopiania_cretica:0.0182831,((Symphytum_caucasicum:5.5836E-4,Symphytum_peregrinum:7.5405E-4):0.00568092,((Symphytum_asperum:0.00158582,Symphytum_tuberosum:0.0037354):9.1348E-4,(Symphytum_ibericum:9.5719E-4,Symphytum_orientale:0.00137503):0.00183302):0.0049486):0.0):0.01723191):0.00726979):9.1518E-4):3.195E-7,(Brunnera_orientalis:0.01319224,((Moritzia_lindenii:0.01721182,(Thaumatocaryon_dasyanthum:0.01182358,Thaumatocaryon_tetraquetrum:0.00700884):0.00702923):0.03252465,(Gastrocotyle_macedonica:0.02065007,((Phyllocara_aucheri:0.01189536,(Anchusa_aegyptiaca:0.00510238,(Anchusa_azurea:0.00461989,Hormuzakia_aggregata:0.00501049):0.00209184):0.00355909):0.00560633,((Anchusa_thessala:0.00873241,((Anchusa_pusilla:1.0E-8,Anchusa_stylosa:0.00130206):0.00464109,(Cynoglottis_barrelieri:0.00377002,Cynoglottis_chetikiana:0.00415684):0.00250697):0.00167258):0.00263943,(Lycopsis_arvensis:0.00163726,Anchusella_variegata:0.01643218,((Anchusa_cespitosa:0.00204729,Anchusa_undulata:4.09E-4):4.09E-4,(Anchusa_crispa:1.0E-8,(Anchusa_capellii:0.00123246,Anchusa_formosa:1.0E-8):5.5344E-4):0.00224589,(Anchusa_capensis:8.6953E-4,Anchusa_leptophylla:0.0015047,Anchusa_officinalis:0.00128534):2.1313E-4):8.8818E-4):0.00431954):0.0098903):5.8894E-4):0.00891088):0.00149564):0.00110732):0.0024243):0.01108685,((Podonosma_orientalis:0.01473354,(Alkanna_orientalis:0.00202902,(Alkanna_pinardii:0.00298669,Alkanna_tinctoria:0.00193309):0.0):0.00708991):0.00902117,((Huynhia_pulchra:0.00861213,(Arnebia_guttata:0.05587543,(Arnebia_benthami:0.00625606,Macrotomia_densiflora:0.02686789):0.00215226):0.00593567):0.00127943,((((Cerinthe_alpina:0.03163343,Cerinthe_major:0.01197551):0.00923484,(Neatostema_apulum:0.01549795,(Moltkiopsis_ciliata:0.03989555,((Lithodora_hispidula:0.00207732,Lithodora_zahnii:0.00178239):0.00771427,(Paramoltkia_doerfleri:0.01416062,(Halacsya_sendtneri:0.01645336,Mairetis_microsperma:0.0244085):6.6314E-4):0.0013828):0.00316376):0.00177625):0.00453931):0.00205894,(('Buglossoides purpureo-caeruleum':0.00732276,(Buglossoides_arvensis:0.0037574,(Buglossoides_tenuiflora:1.0E-8,Buglossoides_incrassata:1.0E-8):0.00831159):0.01134658):8.3711E-4,((Glandora_oleifolia:0.0054778,(Glandora_rosmarinifolia:0.00984785,Glandora_diffusa:0.00516739):0.00594042):0.00302965,(Ulugbekia_tschimganica:0.01545502,((Lithospermum_bejariense:0.0058552,Lithospermum_leonotis:9.2815E-4,Lithospermum_incisum:0.00639745,Lithospermum_macromeria:0.00369854):4.9019E-4,(Lithospermum_distichum:0.00379,Lithospermum_multiflorum:0.00461511,Lithospermum_nelsonii:0.00315846,Lithospermum_officinale:0.00141455):9.9865E-4):0.00205928):0.00324548):0.00137408):0.00622894):8.8059E-4,(((Moltkia_petraea:0.00577212,Moltkia_suffruticosa:0.00730928):0.00467577,(Moltkia_aurea:0.00542395,(Moltkia_angustifolia:0.00979022,Moltkia_caerulea:0.01372934):0.00531492):0.01579102):0.00909167,((Cystostemon_heliocharis:0.01259534,(Maharanga_emodi:0.02131837,(Onosma_alborosea:0.00552505,Onosma_graecum:6.8787E-4,Onosma_taurica:0.00308027,Onosma_visianii:0.0124225,(Onosma_echioides:0.00354325,Onosma_stellulata:0.0036437):0.00122289):0.0152546):0.00222361):0.0042008,((Echium_russicum_Pontechium_maculatum:0.01695039,(Echiostachys_incanus:0.00150726,(Lobostemon_fruticosus_48:0.00378115,Lobostemon_trigonus:0.01127856):0.00213668):0.00290273):9.8951E-4,((Echium_angustifolium:0.00571696,(Echium_italicum:0.00618692,Echium_plantagineum:0.00456304):0.00301057):3.3453E-4,(Echium_vulgare:0.01557268,((Echium_decaisnei:0.00104207,Echium_leucophaeum:8.371E-4):0.00106323,(Echium_aculeatum:0.00172086,(Echium_bonnetii:0.00327419,(Echium_candicans:6.6385E-4,Echium_wildpretii:0.00282192):0.0):0.00113):7.2684E-4):0.00235324):0.00280964):0.00150648):0.01009203):0.00400409):7.9097E-4):4.7755E-4):0.00419309):0.01440791):0.00455784):0.00560554):0.0399087):0.02701794):0.007518095); [! TreeBASE tree URI: http://purl.org/phylo/treebase/phylows/tree/TB2:Tr64871] END;